Glossário - Punt (2007)

Glossary of pollen and spore terminology
⁎ ⁎
W. Punt a, , P.P. Hoen a, , S. Blackmore b, S. Nilsson†, A. Le Thomas c
a Laboratory of Palaeobotany and Palynology, Utrecht University, Utrecht, The Netherlands b Royal Botanic Garden Edinburgh, 20a Inverleith
Row, Edinburgh, EH3 5LP, UK c Laboratoire de Phytomorphologie E.P.H.E., Muséum national d'Histoire naturelle, 16, Rue Buffon, 75005,
Paris, France
Received 28 June 2006; accepted 28 June 2006 Available online 28

November 2006
Abstract
The glossary of pollen and spore terminology was first presented to the international palynological community as the final outcome of
the Working Group on Palynological Terminology at the 8th International Palynological Congress in Aix-en-Provence in 1992. It became widely
accepted as reference guide for palynologists to assist in the preparation of accurate and consistent descriptions of their material. It also serves as a
practical source of information for non-specialists who wish to understand the meaning of the large number of existing palynological terms.
The history of the glossary began in 1972 at the 3rd IPC at Novosibirsk when the working group on palynology was established.
Throughout its history the project has been a collaborative effort with contributions from many palynologists, representing all branches of the
discipline. Only through this long and elaborate procedure, with input from many people, it has been possible to produce the glossary.
The entries are arranged alphabetically and are accompanied by simple schematic illustrations where appropriate. These contain the
minimum amount of information needed to explain the feature. Moreover, to simplify the recognition of pollen and spore wall layers, colours have
been used to indicate the corresponding layers.
The first edition had 547 terms of which 339 have been accepted and recommended for use. In the second edition, a further 41 terms
have been added with their appropriate illustrations. Of these, 10 have been accepted and 31 rejected for a variety of reasons. Where necessary,
illustrations have been revised. An extensive list of consulted literature has been added. © 2006 Elsevier B.V. All rights reserved.
www.elsevier.com/locate/revpalbo
⁎ Corresponding authors.
E-mail addresses: puntw@worldonline.nl (W. Punt), p.hoen@bio.uu.nl (P.P. Hoen).
0034-6667/$ - see front matter © 2006 Elsevier B.V. All rights reserved.
doi:10.1016/j.revpalbo.2006.06.008
Review of Palaeobotany and Palynology 143 (2007) 1–81
2 W. Punt et al. / Review of Palaeobotany and Palynology 143 (2007) 1–81
dicate the different layers more easily. The meaning of the
olours is given in the Introduction.
PREFACE TO THE FIRST EDITION Finally, this second edition is dedicated to the late
r. Siwert Nilssson who, in fact, was the initiator of the first
This Glossary of Pollen and Spore Termi- nolo dition. His continuous support to carry on with the project
is presented to the international palynological community as much helped us to finish the final manuscript.
the occasion of the 5th anniversary of the LPP Foundati Utrecht, December 2005
The publication of the Glossary concludes a project of
Working Group on Palynologi- cal Terminology, function
under the auspices of the International Federation
Palynological Societies (IFPS). Under conve- norship of W
Punt, an international committee of pollen morpholog
has made an up-to-date compilation of the wide diversity
technical terms so far used in the description of pollen a
spores—both modern and fossil. TRODUCTION
Wim Punt, Stephen Blackmore, Siwert Nilsson a

The terminology used in palynology has long been
Annick Le Thomas have to be congratulated with the resu
cognised as a deterrent to those who are not specialists in the subject.
of their effort. Under the technical editorship of Peter Ho
e hope that this glossary will make the subject more widely accessible
who was also responsible for most of the illustrations,
the same time as simplifying the application of palynological terms
format of their Glossary has become fully in harmony w
thout losing any precision. We recognise that the Glossary is not
the twofold objective of the terminology project. Besid
rfect, and anticipate that revisions will be needed in the future.
being a useful reference guide for palynologists who ha
to provide accurate descriptions of their material, This introduction sets out the objectives of the glossary,
Glossary may also serve as a practical source of informatplains the format that has been followed in
for non-specialists who have to understand the meaningext and the illustrations and, for those who may be interested,
an ever-increasing number of palynological terms. rds the history of the project.
Utrecht, December 1993 LPP Foundation,
OBJECTIVES
Henk Visscher (Chairman) Henk
Brinkhuis (Director)
The objective of the project has been to provide a concise
PREFACE TO THE SECOND EDITION
ual of terminology that can be used to clarify the communication of
The first edition of the Glossary of Pollen and mation concerning pollen grains and spores.
Spore Terminology was presented to the internationa It is hoped that this will help to make palynological
palynological community in 1994. During the past 11 years ature more accessible to non-specialists and to beginners in the
has served its purpose well, but during this time severa. In this way it should encourage an increasingly standardised
more terms have been proposed and introduced. Moreoveoach to the description of pollen grains and spores.
a number of mistakes were detected and some drawing We have tried to keep the glossary as simple as possible so
needed adapta- tion. For all these reasons the presen it can easily be used without much previous experience of
authors considered a second, revised edition of the glossarnology.
should be prepared. E FORMAT OF THE GLOSSARY
The setup of the glossary is not changed at all as
has proven its usefulness. We have kept the use o The entries are arranged alphabetically. The form that
indications bold for accepted terms and normal text foears first (the singular, plural or adjectival form of the term) is
rejected terms and synonyms. The indications for plural and erally the most commonly used form, although if all forms are
adjective of the terms has also been main- tained. The mosely used the singular is given first. A number of terms are mainly, or
important difference is the use of colours for the drawingslusively, used as adjectives.
We had in mind that different colours may be useful t Where the term is printed in bold typeface this indicates
that the term is in current usage and is recommended for continuedolid line is used to indicate a feature visible at the surface whilst a
Where the head word is printed in normal text this indicates tha d line indicates a feature that lies below the surface, or behind
alternative term should be used in preference. The definitionser feature.
literature references provided for such terms are included becausetandardised section of a pollen wall, illustrated below (Fig. 1), is
may be helpful in interpreting the literature. In each case the preferas the basis of a number of the diagrams referring to wall
or synonymous term is indicated. fication.
Cross references are also given to terms that indicate
ours are used to indicate the different layers (Fig. 2).
opposite condition (antonyms) and to related terms (indicated by
also”). A comment is provided where this may help in the applicatio
HISTORY OF THE GLOSSARY
a term, or to qualify the circumstances in which it is applied.
The literature reference given for each term is This glossary is the outcome of a initiative that began
necessarily the earliest publication in which the term was used bu the establishment, under the auspices of the International
been selected as a helpful source of further information. mission for Palynology (now, the International Federation of
ological Societies) of a Working Group on Palynological
ILLUSTRATIONS nology. With Jan Muller as Secretary, the Working Group
municated by means of circulars and question- naires distributed
Simple schematic illustrations have been provided wg its members. Siwert Nilsson took on the role of Secretary of the
appropriate. These contain the minimum amount of information neing Group between the 4th International Palynological Congress in
to explain the feature. Some conventions have been used: now (1976) and the 5th IPC held in Cambridge in 1980. A report
progress made by the Group was published by Nilsson and Muller
• Where both equatorial view and polar view are shown, the equat).
view (e) is generally given to the left of the polar view (p). At the 5th IPC it was proposed that the Working Group
• Drawings of pollen grains or spores seen in equatorial view d work towards the publication of a glossary that would
always shown with the distal pole uppermost. arise and explain palynological
terminology. After the
• Features of ornamentation are generally illustrated by a surface viewridge IPC Ste- phen Blackmore became Secretary and
on the left and a sectional view to the right. onnaires continued to be circulated, in
an effort to establish the
aches to terminology that could be adopted in a glossary.
• Arrows have been used to indicate the particular part of the diagra
to which the term applies. In other cases the feature referred tover, as previous experience had shown, relatively few
shaded. In some cases both arrows and shading have been used. ologists replied, calling
Fig. 1. Two different systems used to describe the exine stratification.
debate at the 6th IPC in Calgary (1984), the Working Group was
dissolved with the intention of seeking a more rapid method of
progress. A revitalised Working Group, with Wim Punt as Secretary,
emerged at the 7th IPC in Brisbane (1988). Punt offered to convene a
mall committee that would start working on drafting a glossary. It was
agreed that drafts of the glossary would be circulated as widely as
possible so that the text could be revised as thoroughly as possible
before publication (Report in Palynos 12,2, 1989). To achieve this, it
was decided that draft manuscripts would be circulated to the
representatives
ch of the societies affiliated to the IFPS and to all members of the
y convened Working Group.
Wim Punt then acted as convenor of a committee
isting of Stephen Blackmore, Siwert Nilsson and Annick Le
mas. The First Draft, with a red cover, was circulated in 1989 and
ulated considerable interest. A Second Draft, with an orange cover,
wed in 1990 and again drew many comments and suggestions from
Working Group. These comments were incorporated in a Third
t, not widely distributed, which also included the very detailed
ments of palaeopalynologists Al Traverse and Jan Jansonius.
into question the utility of the Group's methodology. During a

Fig. 2. Colour scheme used in the drawings of the terms.
In April 1991, at the invitation of Knut Fægri the comm g and rather elaborate procedure it has been possible to produce the
met in Bergen to revise the Third Draft and to consider the illustrasent
and publication of the resulting glossary. The process of reconc ary, which we hope goes some way towards meeting the original
out- standing differences of opinion and revising the definitions, staives of the project.
in Bergen, was subsequently continued at meetings in Utrecht
London. NOWLEDGEMENTS
In August 1992 an illustrated version of the Third Draft
prepared by Peter Hoen and distributed at the 8th IPC As in its first edition, the second of the Glossary has
Aix-en-Provence in a pale yellow cover as the First Concept. O benefited greatly from the advise and expertise of a number of
again the project benefited from the detailed comments of m ologists. We are in particular grateful for the remarks and critical
palynologists. These have been included in this edition. ment of Dr. J. Jansonius and Dr. M. Harley.
It will be clear, from the history of the project, that thi We thank the technical staff of the Elsevier Geoscience
been a collaborative project, with contributions from m als Department for their help and continuous encouraging support
palynologists, from all branches of the discipline. Only throughking the Glossary project a success.
Last but not least we would like to thank Prof. Dr. ology for their everlasting support of this project.
Lotter and the members of the Laboratory of Palaeobo- tany a
THE GLOSSARY
In all drawings, the distal pole is on top, the proximal pole at the bottom, unless otherwise specified. e = equatorial view, p
= polar view; H = high level, L = low level. The terms printed in bold are preferred.
A-, an-
A prefix indicating the absence of a feature, as for example in alete.
A-type tetrad (Moar, 1993)

A tetrad in which the aborted cells (1–3 in number) are clearly associated with
the fertile grain(s). Example: Cyathodes juniperina (Epacridaceae). See also: S-type tetrad,
T-type tetrad.
Abporal lacuna (pl. abporal lacunae) (Wodehouse, 1928)

A lacuna in a lophate pollen grain situated at the end of an ectoaperture that is
divided by sexinous ridges into two or more lacunae. Example: Sonchus oleraceus
(Compositae). See also: lacuna, lophate.
Acalymmate (adj.) (Van Campo and Guinet, 1961)

Describing tetrads or polyads in which the sexine/ectexine of each monad is well
differentiated, but does not form a single continuous envelope around the unit. Example:
Drosera (Droseraceae). Antonym: calymmate.
Acanthomamilla (Balme, 1988)

A biform sculptural element consisting of a hemispheroidal base, surmounted by
a sharply contracted spine. Examples: Acinosporites, Dibolisporites.
Acetolysis (Erdtman, 1960a)

A widely used technique for preparing pollen and spore exines for study.
Acolpate (adj.) (Moar, 1993)

Without colpi.
Acrolamella (pl. acrolamellae, adj. acrolamellate) (Li and Batten, 1986)

A leaf-like tapering segment on the proximal pole of a megaspore. Example:
Arcellites. Comment: An acrolamella is a special form of gula, represented by leaf-like
segments. See also: gula, trifolium.
Actuopalynology (Hulshof and Manten, 1971)
The study of pollen grains and spores of extant plants.
Antonym: palaeopalynology. See also: pollen analysis.
Aeropalynology (Erdtman, 1969)
The study of palynomorphs found in the atmosphere.
-alA suffix for of, relating to, or characterized by. Example: tectal.
Alete (adj.) (Erdtman, 1943)

Describing a spore without a laesura. See
also: -lete, laesura.
Alveolate (adj.) (Van Campo, 1971)

Describing a type of sexine/ectexine structure, in which the infratectal layer is
characterised by partitions forming compartments of irregular size and shape. Example:
Pinus (Pinaceae). See also: columella, granular exine.
Amb (Erdtman, 1952)

The outline of a pollen grain or spore seen in polar view. Comment: The term
does not necessarily coincide with the equatorial outline. The term is especially useful
for the outline of spores seen in polar view, because of their heteropolar shape.
Ana-(Erdtman and Vishnu-Mittre, 1956)

A prefix indicating the position of features, such as apertures, on the distal
face. Examples: Nymphaea (Nymphaeaceae), Sparganium (Typhaceae). Antonym: cata-.
See also: zona-.
Anazonasulculate (adj.) (Walker and Doyle, 1975)

Describing a pollen grain with a ring-like sulculus situated between the
equator and the distal pole. Antonym: catazonasulculate.
Ancyrate (adj.) (Balme, 1988)

Bearing sub-cylindrical or tapering processes which divide at their distal
extremities into anchor-shaped or multifurcate tips.
Angulaperturate (adj.) (Erdtman, 1952)

Describing an equatorially aperturate pollen grain with the apertures situated at
the angles of the outline in polar view. Example: Corylus (Betulaceae). Antonym:
planaperturate. Comment: The term is useful for describing the position of apertures, but
should be avoided as a description of equatorial outline. See also: sinu-aperturate,
fossaperturate.
Adequate (adj.) (Reitsma, 1970)
Synonym of spheroidal.
Angustimurate (adj.) (Erdtman, 1952)
With narrow muri. Comment: Erdtman restricted the term to muri up
to 1/5th of the diameter of the lumina.
Anisopolar (adj.) (Erdtman, 1947)

Synonym of heteropolar.
Anisodiametric tetrad
General term for tetrads which members are differing in size. Example:
seed-megaspore tetrads. Antonym: isodiametric tetrad.
Annulus (pl. annuli, adj. annulate)

An area of the exine surrounding a pore that is sharply differentiated from the
remainder of the exine, either in ornamentation or thickness (Jackson, 1928). Comment:
Although the correct Latin spelling is anulus, the form used throughout the botanical
literature is annulus. See also: aspis, costa, margo.
Anulus (Beug, 1961)
Orthographical variant of annulus.
Anteturma (pl. anteturmae) (Potonié, 1956)

An artificial grouping of fossil spores and pollen in the turma-system of Potonié.
Comment: Turmae are grouped under two large headings the anteturmae Sporites and
Pollenites. See also: infraturma, subturma, turma.
Aperture (adj. aperturate) (Erdtman, 1947)

A specialized region of the (sporoderm, that is thinner than the remainder of the
sporoderm and generally differs in ornamentation and/or in structure. Comment: Apertures
are described as simple if they are present in only one wall layer, or compound if they affect
more than one layer of the wall. In compound apertures the shape of apertures may differ
between layers. Ectoapertures occur in the sexine/ectexine, endoapertures in the
nexine/ endexine, andmesoaperturesare sometimes found in an intermediate position
between an ecto- and endoaperture. Various types of apertures are recognised on the basis
of their shape (see, colpus, laesura, porus, sulcus, ulcus), position (see, ana-, cata-,
zona-, zono-) or fusion (see, syn-). In living pollen grains or spores the apertures usually
function as sites of germination, they may also provide routes for transfer of water and
other substances, and play a part in harmomegathy. The term is often used in
conjunction with a prefix or suffix, as for example in, ectoaperture, endoaperture,
inaperturate, omniaperturate, pseudoaperture, triaperturate.
Apex (pl. apices)
A general term for the tip of an organ (Jackson, 1928). Comment: In fossil
spores applied to the tip or corner of a trilete spore.
Apicalfeld (Beug, 1961)

Synonym of apocolpial field.
Apiculate (adj.) (Fægri and Iversen, 1950)

Describing the shape of a pollen grain that has slightly protruding polar caps.
Example: Adonis aestivalis (Ranunculaceae).
Apiculate elements (Smith and Butterworth, 1967)

Projections from the general surface. Examples: bacula, pila,
verrucae, spines.
Apocolpial field (Punt et al., 1974)

A region at the pole of a parasyncolpate pollen grain, delimited by the
margins of the anastomosing colpi. See also: apocolpium.
Apocolpium (pl. apocolpia) (Erdtman, 1952)

A region at the pole of a zonocolpate pollen grain delimited by lines connecting
the apices of the colpi. Synonym of polar area. Comment: In the original definition the
apocolpium was delimited by the polar limits of the mesocolpia, the more precise definition
now used is derived from Iversen and Troels-Smith (1950, polar area). See also:
apoporium, mesocolpium.
Apocolpium index (Punt, 1976)

The ratio of the distance between the apices of two ectocolpi (d) of a
zonocolpate pollen grain to its equatorial diameter (D). Synonym of polar area index.
Aperture membrane (Erdtman, 1952)
The exine which forms the floor of an ectoaperture. Comment: Depending on
the type of ectoaperture these may be colpus membranes or pore membranes,
usually representing the nexine. The membrane may be partially or completely covered by
an operculum.
Arcus (pl. arcus, adj. arcuate) (Erdtman, 1947)

A locally thickened band of sexine that extends in a sweeping curve from one
aperture to another. Example: Alnus (Betulaceae). Comment: In palynological literature the
plural is often misspelled as arci. However the correct plural is arcus.
Areola (pl. areolae, adj. areolate) (Erdtman, 1947)

A feature of ornamentation in which the sexine/ectexine is composed of circular
or polygonal areas separated by grooves which form a negative reticulum. Examples:
Apama (Aristolochiaceae), Phyllanthus spp. (Euphorbiaceae).
Aspidote (adj.) (Erdtman, 1952)

An orthographical variant of aspidate. See
also: aspis.
Aspis (pl. aspides, adj. aspidate) (Wodehouse, 1935)

A prominently protruding thickening of the exine around a pore. Examples:
Betula (Betulaceae), Dorstenia (Moraceae). Comment: An aspis is a special form of
annulus, represented by a thickening rather than a thinning. See also: atrium.
Aspis channel (Engel, 1980)

Synonym of pore canal.
Apolar (adj.) (Erdtman, 1952)
Describing pollen and spores without distinct polarity. Example:
Plantago (Plantaginaceae).
Apoporium (pl. apoporia) (Erdtman, 1952)

An area at the pole of a zonoporate pollen grain that is delimited by a line
connecting the borders of the pores. Comment: The term contrasts with apocolpium and is
most meaningful in pollen grains with large pores. See also: apocolpium, mesoporium.
Auricula (pl. auriculae, adj. auriculate) (Potonié and Kremp, 1955)

Synonym of valva. Comment: The term is mostly used for spores with
extreme valva.
Baculum (pl. bacula, adj. baculate) (Potonié, 1934)

A cylindrical, free standing exine element more than 1 μm in length and less
than this in diameter. Example: Raistrickia saetosa. See also: columella, pilum.
Bilateral (adj.)
Describing pollen and spores having a single, principal plane of symmetry.
Comment: According to Walker and Doyle (1975) bilaterally symmetrical objects could
more correctly described as anisobisymmetric, but because of the familiarity of the term
bilateral it was used by them.
Bireticulate (adj.) (Bor, 1979)

A two-layered reticulum consisting of a suprareticulum supported by a
microreticulate tectum. Examples: Entelea arborescens (Tiliaceae), Phyllanthus
oppositifolius (Euphorbiaceae), Salvia azurea (Lamiaceae).
-ateA suffix for possession of. Example: porate.
Atectate (adj.) (Walker and Doyle, 1975)

Describing pollen grains that have an exine with little or no internal structure.
Example: Degeneria (Degeneriaceae). Comment: The term was originally intended to
describe walls of primitive angiosperms lacking columellae. Related terms include
intectate, which is used for pollen grains with columellae but without a tectum, and
etectate which is applied to pollen grains believed to have lost their tectum during
phylogeny. See also: etectate, intectate, tectum.
Atrium (Thomson and Pflug, 1953)

A space within the aperture of a compound pore that has a much larger
endopore than the ectopore, so that the pore canal widens towards the interior of the
grain. Example: Myrica (Myricaceae). See also: vestibulum.
Atrium (Punt, 1962)

Synonym of fastigium.
Bisaccate (adj.) (Potonié and Kremp, 1954)

Describing pollen with two sacci. Example: Pinus (Pinaceae). See
also: saccus, protosaccus, pseudosaccus.
Brachy-
A prefix for short.
Synonym of brevi-.
Brevi-
A prefix for short. Example: brevicolpate.
Breviaxe (adj. breviaxal) (Van Campo, 1966)
Pollen grains with a polar axis that is shorter than their equatorial diameter.
Synonym of oblate (s.l.). Antonym: longiaxe. Comment: Thomson and Pflug (1953)
recognised Brevaxones a group of mid-Cretaceous and later angiosperm pollen grains
that was contrasted with Longaxones. See also: successiform.
Breviaxy (Van Campo, 1966)

A phyletic series of pollen forms ranging from subspheroidal tricolpate, to
tricolporate, to oblate tricolporate. See also: successiform.
Brevissimi-
A prefix for very short.
Bridge (Fægri and Iversen, 1950)

A feature in which the margins of the colpi are raised in the equatorial region and
connected with each other, forming a bridge over the ectocolpus and dividing it into two
parts. Example: Hemandradenia (Connaraceae). See also: demicolpus.
Brochus (pl. brochi, adj. brochate) (Erdtman, 1952)

A brochus consists of one lumen of a reticulum and half of the width of the
adjacent muri. See also: heterobrochate, homobrochate, lumen.
Callose wall (Mangin, 1899) Part of the

special wall.
Calymmate (adj.) (Van Campo and Guinet, 1961)

Describing tetrads or polyads in which the sexine/ectexine of each monad is well
differentiated and forms a continuous envelope around the unit. Example: Acacia
(Mimosaceae). Antonym: acalymmate.
Caput (pl. capita, adj. capitate) (Erdtman, 1952)
The expanded apex (head) of a columella. See
also: columella, pilum.
Cata- (Erdtman and Vishnu-Mittre, 1956)

A prefix indicating the location of features, such as apertures, on the proximal
face. Antonym: ana-. See also: zona-.
Camera (pl. camerae, adj. camerate) (Neves and Owens, 1966)
A cavity formed by the separation of two wall layers in spores that lacks an
infrastructure. See also: cavus, pseudosaccus, saccus.
Canaliculate (adj.) (Potonié, 1934)
Synonym of fossulate.
Capillus (pl. capilli, adj. capillate) (Potonié and Kremp, 1955)

Synonym of fimbria.
Cappa (pl. cappae) (Erdtman, 1957)

The thick-walled proximal side of the corpus of a saccate pollen grain.
Cappula (pl. cappulae) (Erdtman, 1957)

The thin-walled distal side of the corpus of a saccate pollen grain.
Synonym of leptoma.
Capsula (pl. capsulae, adj. capsulate) (Pocock, 1961a)

An outer structure of a spore projecting at the equator and completely enclosing
the spore body. See also: cingulum, patina.
Cicatricose (adj.) (Potonié, 1934)
Describing spores marked with scars. Example: Cicatricosisporites.
Catazonasulculate (adj.) (Walker and Doyle, 1975)
Describing a pollen grain with a ring-like sulculus situated between the equator
and the proximal pole. Antonym: anazonasulculate.
Cavate (adj.)
In spore terminology, a synonym for camerate. See also:
camera.
Cavea (pl. caveae, adj. caveate) (Skvarla and Larson, 1965)

A cavity between two layers of the exine extending to the colpus margin where
the layers meet. Example: Ambrosia (Compositae).
Caverna (pl. cavernae) (Thomson and Pflug, 1953)

Synonym of fastigium. Comment: Originally a distinction was made between a
separation of two layers of the ectexine, which was called a praecaverna, and a separation
within the endexine, a postcaverna. The term is not used to describe recent pollen grains.
Cavium (Thomson and Pflug, 1953)

A subpolar chamber formed at the end of three anastomosing cavernae.
Cavum (pl. cava, adj. cavate) (Fægri and Iversen, 1989)

Synonym of cavea.
Cavus (sensu Skvarla and Turner, 1966)

Synonym of cavea.
Centrosymmetrical (adj.) (Straka, 1964)

Synonym of radially symmetric.
Cingulum (pl. cingula, adj. cingulate) (Potonié and Kremp, 1955)
A thick outer structure of a spore that projects at the equator, but does not
extend over the distal or proximal face. Example: Densosporites anulatus. See also:
capsula, patina.
Circumaperturate (adj.) (Straka, 1964)

Describing a pollen grain with equatorial apertures that are regularly arranged
around a circular outline.
Circumpolar lacuna (pl. circumpolar lacunae) (Wodehouse, 1928)
A lacuna in lophate pollen grain immediately adjacent to a polar lacuna that is
not part of the apertural system. See also: lacuna, lophate.
Clava (pl. clavae, adj. clavate) (Iversen and Troels-Smith, 1950)

A club-shaped element of the sexine/ectexine that is higher than 1 μm, with
diameter smaller than height and thicker at the apex than the base. See also: baculum,
columella, gemma, pilum.
Coaperturate (adj.) (Beug, 1961)

Describing permanent tetrads in which the apertures of neighbouring monads
join. Examples: Erica (Ericaceae), Periploca (Periplocaceae). See also:
syncolp(or)ate, Fischer's law/rule.
Colpodiporate (adj.) (Selling, 1947)

Synonym of diploporate.
Colpororate (adj.) (Moar, 1993)

A compound aperture characterised by an ectoaperture, a shorter lolongate
mesoaperture and a lalongate endoaperture. Example: Sonchus (Compositae). Comment:
This term descibes a compound aperture with a mesoaperture. See Also: mesoaperture.
Colpoid (Erdtman, 1952)

Apertures more or less similar to colpi but less clearly defined in outline.
Colporoidate (adj.) (Erdtman, 1952)

Describing pollen grains with colpi and indistinct ora.
Colpus (pl. colpi, adj. colpate) (Erdtman, 1943)

An elongated, aperture with a length/breadth ratio greater than 2. Comment:
Some authors restrict the term to a meridional aperture (and contrast this with a distal or
proximal sulcus), but since shape is the defining criterion the term is equally applicable to
other locations (for example, pantocolpate pollen). The precise meaning is often
indicated by the use of a prefix, as for example in, ectocolpus, endocolpus,
syncolpate.
Colpus equatorialis (pl. colpi equatoriales) (Iversen and Troels-Smith, 1950)

Synonym of endocingulum.
Colpus membrane (Iversen and Troels-Smith, 1950)

The aperture membrane of a colpus.
Colpus transversalis (pl. colpi transversales) (Wodehouse, 1935; Fægri and

Iversen, 1950)
Synonym of endocolpus. See
also: endoaperture.
Columella (pl. columellae, adj. columellate) (Iversen and

Troels-Smith, 1950)
A rod-like element of the sexine/ectexine, either supporting a tectum or a
caput. Comment: The difference between a baculum and a columella in current usage is,
that a baculum is always a free standing element of sculpturing, whereas a columella is part
of the structure. See also: baculum, pilum.
Columellae layer (Reitsma, 1970)

Synonym of infratectum. See
also: interstitium.
Colporus (pl. colpori, adj. colporate) (Erdtman, 1945a)
A compound aperture consisting of an ectocolpus with one or more
endoapertures. Comment: The term is most commonly used in its adjectival form,
colporate.
Compound aperture (Erdtman, 1969)
An aperture with two or more components that are situated in more than one
wall layer.
Concordant pattern (Fægri and Iversen, 1989)

A pattern in a tectate pollen grain in which the arrangement of the columellae is
the same as that of the elements upon the tectum. Example: Lilium (Liliaceae). Antonym:
discordant pattern.
Conjunctate (adj.) (Skvarla and Larson, 1965)

With bacula or columellae which are branched proximally into two or more
parts.
Contact area (Potonié, 1934)

Area on the proximal face of a spore interpreted as having been formed in
contact with the other members of the tetrad. Example: Retusotriletes pythovii.
Conus (pl. coni) (Potonié and Kremp, 1955)

Cone-shaped elements on the surface of spores in which the height is less than
two times the basal diameter and the apex is pointed, blunt or rounded. Example:
Lophotriletes mosaicus. Comment: The term is only used in spore terminology and is more
or less synonymous with spine.
Copropalynology (Erdtman, 1969)

The study of palynomorphs in coprolites or excrement.
Commissure (Harris, 1955)
The slit or line of dehiscence in the laesura. See
also: laesura.
Composite aperture (Erdtman, 1952) Synonym of

compound aperture.
Costa equatorialis (pl. costae equatoriales) (Iversen and Troels-Smith, 1950)

Synonym of costa endocinguli.
Crassi-
A prefix for thick.
Crassitude (Grebe, 1971)

Synonym of a thickening.
Crescentic (adj.) (Fægri and Iversen, 1989)

Describing a pollen grain in polar view with a very thick exine in the medium of
the intercolpium, gradually thinning towards the colpi. Example: Valerianella
(Valerianaceae).
Crista (pl. cristae, adj. cristate) (Potonié and Kremp, 1955)
A crest-like ornamentation element, taller than it is wide, characterized by a
narrowly curved base and a sharp upper edge. Example: Cristatisporites.
Cross tetrad (Erdtman, 1945b)

Synonym of decussate tetrad.
Corona (pl. coronae, adj. coronate) (Potonié and Kremp, 1955)
An equatorial or subequatorial extension of a spore, resembling a
cingulum, but divided into fringe-like elements (fimbria). Example: Reinschospora
speciosa.
Corpus (pl. corpi) (Erdtman, 1957)

The body of a saccate pollen grain or camerate spore.
Corrugate (adj.) (Erdtman, 1947)

Synonym of rugulate.
Costa (pl. costae, adj. costate) (Iversen and Troels-Smith, 1950)

A thickening of the nexine/endexine bordering an endoaperture, or following the
outline of an ectoaperture. Comment: The following forms are used: costa ectocolpi,
costa ectopori, costa endocinguli, costa endocolpi, costa endopori.
Cuneus (pl. cunei) (Batten and Christopher, 1981)
A structure formed by the termination of the inner wall layer at the
endogerminal which gives rise to a shallow V-shaped atrium that points towards the centre
of the grain. Example: Pseudotrudopollis. Comment: This term is used in descriptions of
the fossil Normapolles group. See also: incidence.
Cuniculus (pl. cuniculi) (Sullivan, 1964)

A space between the margin of the spore body and the inner face of the
equatorially expanded spore wall, located at the equator. See also: camera.
Cupulate (Gupta and Udar, 1986)

Synonym of foveolate.
Croton pattern (Erdtman, 1952)
A characteristic type of ornamentation comprising rings of five or six
(sometimes more) raised, often triangular, sexine elements arranged around a circular area,
usually formed by capitate columellae (pila). Example: Croton, Jatropha (Euphorbiaceae),
Callitriche antartica (Callitrichaceae), Pimelea arenaria (Thymelaceae).
Crustate (adj.) (Erdtman, 1952)

Describing an aperture membrane that is thickly covered with coarse
granules.
Cryptoaperture (adj. cryptoaperturate) (Thanikaimoni, 1980)

An endoaperture which is not apparent in surface view, because there is no
ectoaperture. Comment: Examples of such apertures have been described in Phaleria
(Thymelaeaceae) (Erdtman, 1952), Tetracera (Dilleniaceae) (Kubitzki and Baretta-Kuipers,
1969) and Phyllanthus (Euphorbiaceae) (Meewis and Punt, 1983).
Cryptopolar (Gupta and Udar, 1986)

Describing a spore in which the distal and proximal faces have dissimilar
sculpturing and which lacks tetrad mark. Example: Calobryum dentatum, Haplomitrium
hookeri.
Cryptospore (Richardson et al., 1984; Strother, 1991)

Non-marine alete sporomorph with well differentiated contact areas but
without (haptotypic) features such as laesurae.
Curvatura (pl. curvaturae) (Potonié, 1934)
A line in trilete spores, extending from the extremities of the ends of the radii of
the laesura and thus delimiting the contact areas. Examples: Laevigatisporites glabratus,
Divisisporites divisus. Comment: Curvaturae perfectae are continuous around the
proximal face whereas curvaturae imperfectae are present as forked extensions that do
not join.
Curvimurate (adj.) (Erdtman, 1952)

Describing pollen grains with curved muri.
Decussate tetrad (Walker and Doyle, 1975)

A multiplanar tetrad of pollen grains or spores arranged in two pairs lying across
one another, the pairs (dyads) more or less at right angles to each other. Example: Orophea
(Annonaceae).
Demicolpus (pl. demicolpi, adj. demicolpate) (Erdtman, 1952) An ectocolpus

divided into two parts. Example: Amylotheca (Loranthaceae). See also: bridge.
Di-A prefix for two. Comment: Examples: dicolpate, dicolporate, diporate,

disulcate, disulculate, diulculate.
Dicolpate, dicolporate, diporate (adj.) (Iversen and Troels-Smith, 1950)

Describing pollen grains with two ectocolpi, two compound apertures or two
pores. Comment: The terms belong to the system of pollen classes introduced by Iversen
and Troels-Smith. See also: disulcate, monoaperturate, pantoaperturate,
tricolpate, zonoaper-urate.
Dicolporate (adj.) (Cranwell, 1953)

Synonym of diploporate. Comment: The same term refers to a pollen class in
the system of Fægri and Iversen (1950).
Digitate (adj.) (Skvarla and Larson, 1965)
With bacula or columellae which are branched distally into two or more
parts. Example: Polygonum bistorta (Polygonaceae).
Diorate (adj.) (Erdtman, 1952) Synonym of

diploporate.
Diploporate (adj.) (Fægri and Iversen, 1964)

Describing an ectocolpus with two endoapertures. Example: Didymeles
(Didymelaceae). See also: endoaperture area.
Diploxylonoid (adj.) (Traverse, 1988)

Describing bisaccate pollen grains in which the outline of the sacci in polar view
is discontinuous with the outline of the corpus so that the grains seem to consist of three
distinct, more or less oval parts. See also: haploxylonoid, Haploxylon-type,
Sylvestris-type.
Discordant pattern (Fægri and Iversen, 1989)

A pattern in a tectate pollen grain in which the arrangement of the columellae is
different from that of the elements on the tectum. Example: Geranium (Geraniaceae).
Antonym: concordant pattern.
Dispersal unit
The morphological unit in which mature pollen grains or spores are shed, which
may range from individuals (monads), to pairs (dyads), groups of four (tetrads), or
groups of more than four (polyads). Larger, indeterminate numbers of pollen grains or
spores may also be dispersed as pollinia or massulae.
Dissections (Couper and Grebe, 1961)

Rounded to elongated cavities in a cingulum or zona. Example: Vallatisporites
ciliaris. Synonym of vacuoles.
Distal (adj.)
A common descriptive term (Jackson, 1928) used in contrast to proximal,
applied in palynology to features on the surface that face outward in the tetrad stage
(Wodehouse, 1935). Antonym: proximal. See also: ana-. cata-, polarity.
Distal face (Erdtman, 1952)

That part of a palynomorph that faces outwards the centre of the tetrad, between
equator and distal pole. Antonym: proximal face.
Distal pole (Erdtman, 1952)

The centre of the surface of the distal face.
Antonym: proximal pole. See also: distal.
Disulcate (adj.) (Selling, 1947; Harley, 1998)

Describing pollen grains with sulci arranged in pairs. Two types of disulcate
pollen are distinguished: equatorial disulcate, with opposing, equatorially arranged
sulci and distal disulcate with paired sulci lying parallel to the long axis of the pollen
grain on the distal face. Examples: Metroxylon salomonense (Palmae) (equatorial
disulcate), Chamaerops humilis (Palmae) (distal disulcate). See also: dicolpate,
dicolporate, diporate, geminicolpate.
Duplibaculate (adj.) (Erdtman, 1952) Synonym of

duplicolumellate.
Duplicolumellate (adj.) (Reitsma, 1970)
With columellae in two rows under each murus. Example: Polygonum
persicaria (Polygonaceae). See also: pluricolumellate, simplicolumellate.
DyadGeneral term for two microspores (pollen grains or spores) united as a dispersal unit.
Comment: A developmental dyad stage occurs prior to tetrad formation in plants with
successive meiosis. See also: dispersal unit.
Echinolophate (adj.) (Wodehouse, 1928)

Describing a lophate pollen grain with echinate ridges. See
also: fenestrate, psilolophate.
Ectexine (sensu Fægri, 1956)

The outer part of the exine, which stains positively with basic fuchsin in optical
microscopy and has higher electron density in conventionally prepared TEM sections.
Orthographical variant: ektexine. Comment: Ectexine includes the foot layer (nexine 1), if
present. Erdtman introduced the term in 1943, but used sexine in his later publications. See
also: sexine, endexine.
Ecto-
A prefix for outer.
Ectoaperture (Van Campo, 1958)

An aperture in the outer layer of the sporoderm.
Comment: Examples: ectocolpus, ectopore. See also:
endoaperture, mesoaperture.
Ectointine (Freytag, 1968) Synonym of
exintine.
Ektannulus (pl. ektannuli) (Batten and Christopher, 1981)

An abrupt thickening of the outer wall layer in the region of the exogerminal.
Example: Krutzschipollis. Comment: This term is used in descriptions of the fossil
Normapolles group for part of an annulus.
Ektexine (Erdtman, 1943)

Orthographic variant of ectexine. Comment: The original spelling was soon
superseded by the latinized spelling “ectexine” in the publications of Erdtman and others.
Ekto-Orthographical variant of the prefix ecto-.

Echinate (adj.) (sing. echinus, pl. echinae) (Wodehouse, 1928)
Describing pollen and spores with an ornamentation comprising spines longer
than 1 μm. Comment: Although the correct Latin spelling for the plural is echini, the form
used throughout the palynological literature is echinae. Comment: (Erdtman (1952)
recommended the term spinose, but in his usage spines were defined as longer than 3 μm
and smaller features as spinules (adj. spinulose). See also: microechinate, spine,
spinule.
Endannulus (pl. endannuli) (Thomson and Pflug, 1953)

An annulus formed by the endexine of a pollen grain. Example: Atlantopollis.
Comment: This term is used in description of the fossil Normapolles group.
Endexine (sensu Fægri, 1956)

The inner part of the exine which remains relatively unstained with basic fuchsin
in optical microscopy and has a lower electron density in conventionally prepared TEM
sections. Synonym of nexine 2 (Erdtman, 1960b). Comment: Originally endexine was
defined simply as the inner of the two main layers of the exine (Erdtman, 1943) to which
Erdtman later applied the term nexine. However, as more recently defined, endexine does
not include the foot layer (nexine 1) which is considered part of the ectexine. See also:
ectexine.
Endintine (Kress and Stone, 1982)

The inner, cellulosic zone of the intine which is adjacent to the cytoplasm and in
fluorescence microscopy stains positively with PAS calcofluor. Antonym: exintine.
Orthographical variant: endointine.
Endo-
A prefix for inner.
Endoaperture (Van Campo, 1958)

An aperture in the inner layer of the sporoderm, often the inner aperture of a
compound aperture. Comment: Examples are: endocingulum, endocolpus,
endopore. See also: ectoaperture, mesoaperture.
Endoaperture area (Verbeek-Reuvers, 1976)

The region of the nexine of a single compound aperture which encloses a number
of endoapertures. Comment: The term is used where the number of endoapertures is more
than two and also where the endoapertures are not sharply delimited. Examples:
Deplanchea (Bignoniaceae), Phyllanthus spp. (Euphorbiaceae). See also: colpodiporate,
diploporate.
Endocingulum (pl. endocingula, adj. endocingulate) (Reitsma, 1966)

A ring-shaped endoaperture continuous around a pollen grain and lying in the
equatorial plane. Example: Polygonum convolvulus (Polygonaceae). Synonym of colpus
equatorialis and zonorate.
Elater
A spirally twisted, filamentous band attached to certain spores and some fossil
pollen grains (Jackson, 1928). Example: Equisetum (Equisetaceae).
Endocrack (Oldfield, 1959)
An irregular groove occurring in the inner surface of the nexine/endexine and
readily apparent in acetolysed pollen. Example: Anemone nemorosa (Ranunculaceae).
Endogerminal (adj.) (Batten and Christopher, 1981) Describing an aperture in the inner
wall layer. Comment: This term, used in descriptions of the fossil Normapolles group, is
essentially synonymous with endoaperture.
Endointine (Freytag, 1968)

Orthographic variant of endintine.
Endoplica (pl. endoplicae) (Thomson and Pflug, 1953)

A fold or swelling of the inner wall layer. Comment: A term originally used for
features in fossil pollen of the genus Triatriopollenites, and later used in the fossil
Normapolles group where endoplicae are present as Y-shaped structures centred over the
poles.
Endosculpture (Van Campo, 1971)
Sculpturing occurring on the inner surface of the nexine/endexine.
Endospore
The innermost layer of a spore wall (Jackson, 1928). Comment: It is probably
homologous with the intine of a pollen grain. Also used as a synonym for the corpus in
camerate spores. Mycologists use the term for spores formed on the inside of a
sporangium. See also: exospore, perispore.
Endosporium (Erdtman, 1943)

Synonym of intine.
Equator (Wodehouse, 1935)

The dividing line between the distal and proximal faces of a pollen grain or
spore. See also: equatorial diameter, equatorial plane.
Equatorial axis
Often misappropriately used as a synonym of equatorial diameter.
Equatorial lacuna (pl. equatorial lacunae) (Wodehouse, 1928)

A lacuna situated on the equator in the mesocolpial region of a lophate pollen
grain. Example: Tragopogon pratensis (Compositae). See also: lacuna.
Equatorial limb
Synonym of equatorial outline.
Equatorial outline
General description of the equator when a pollen grain is seen in polar view.
Synonym of limb. See also: amb.
Equatorial plane (Fægri and Iversen, 1950)

The plane perpendicular to the polar axis and lying midway between the poles.
Equatorial ridge (Wodehouse, 1928)

A ridge lying along the equator in the mesocolpial region, usually applied to
lophate pollen grains. Example: Cichorium intybus (Compositae).
Equatorial bridge (Moore and Webb, 1978)
Synonym of bridge.
Equatorial diameter (Erdtman, 1943)

A line, lying in the equatorial plane, perpendicular to the polar axis and passing
through it. Comment: In bilateral palynomorphs the longest line is usually taken to be the
equatorial diameter.
Eu-A prefix for true. Comment: When applied to patterns of structuring, such as
eurugulate, eustriate and eureticulate, the prefix indicates the arrangement of the
columellae. This contrasts with patterns on the tectum which are indicated by the prefix
supra-. See also: supra-.
Euintine (Kuprianova, 1948) Synonym of

endintine.
Eurypalynous (adj.) (Erdtman, 1952)

Describing plant taxa characterized by possession of a great diversity of
palynomorphs. Antonym: stenopalynous.
Eutectum (adj. eutectate)

Describing a pollen grain with a continuous tectum. Synonym of
pertectate, tectum imperforatum and tectum solidum. Antonym: semitectum.
See also: tectum.
Equatorial view (Erdtman, 1943)
The view of a pollen grain or spore where the equatorial plane is directed
towards the observer. Antonym: polar view.
Equiaxe (adj.equiaxal) (Van Campo, 1966)
Pollen grains with a polar axis equal to the equatorial diameter. See
also: breviaxe, longiaxe.
Erect (adj.) (Reitsma, 1970)

Synonym of prolate.
Etectate (adj.) (Walker, 1976)

Describing pollen grains interpreted as having lost their tectum during
evolutionary development. See also: atectate, intectate, tectum.
Exospore
The outer layer of a spore wall (Jackson, 1928). Comment: A perispore, if
present, lies outside the exospore, and is not part of it. Probably homologous with exine in
pollen. Mycologists use the term for spores formed on the outside of a sporangium. See
also: endospore, perispore.
Exine (pl. exines, adj. exinal, exinous) (Fritzsche, 1837)
The outer layer of the wall of a palynomorph, which is highly resistant to
strong acids and bases, and is composed primarily of sporopollenin.
Exine 1 (Erdtman, 1969) Synonym of
ectexine.
Exine 2 (Erdtman, 1969) Synonym of

endexine.
Exintine (Kress and Stone, 1982)

An outer (pectic) layer of the intine, lying below the nexine, and which stains
positively with alcian blue. Synonym of ectointine. Antonym: endintine.
Exitus (Wodehouse, 1935)

The site of exit of the pollen tube from the sporoderm. See also:
aperture.
Exoexine (Potonié, 1934) Synonym of

sexine.
Exogerminal (adj.) (Batten and Christopher, 1981)

Describing an aperture formed in the outer wall layer of the exine. Comment:
This term, used in descriptions of pollen of the fossil Normapolles group, is essentially
equivalent to ectoaperture.
Exosporium (Erdtman, 1943)
Synonym of exine.
Fastigium (pl. fastigia, adj. fastigiate) (Reitsma, 1966)

Cavity in a colporate grain, appearing as a separation of the inner part of the
exine from the domed sexine in the region of the endoaperture. Example: Hypericum
(Hypericaceae). Synonym of caverna. Comment: The term describes a feature
morphologically similar to the vestibulum; the latter is restricted to porate pollen grains.
See also: vestibulum.
Fenestrate (adj.) (Iversen and Troels-Smith, 1950)
Describing a class of pollen grains characterized by large, window-like spaces
lacking a tectum. Comment: The term is accepted as a category in the classification system
of Iversen and Troels-Smith (1950) and was especially established to include lophate pollen
grains of the Compositae. Although useful for defining a class of pollen grains the term is
not recommended in descriptions. Many such grains can be described as lophate.
Fimbria (pl. fimbriae, adj. fimbriate)

Long, hair-like appendages (Jackson, 1928). Comment: The term has been
used in the descriptions of fossil spores. Example: Radiatisporites radiatus. Synonym of
capillus.
Fischer's law/rule (Erdtman, 1952)

The name given to the widespread arrangement in developmental tetrads of
tri-aperturate pollen grains whereby the apertures form in pairs at six points in the tetrad.
Example: Ericaceae. See also: Garside's law/rule.
Fissura (pl. fissurae) (Potonié, 1934)

A sharp, straight split that appears during germination in certain inaperturate
pollen grains. Examples: Taxodium (Taxodiaceae), Cupressus (Cupressaceae).
Flange
A general term, used to describe equatorial extensions of spores (Jackson,
1928). Comment: This term is widely used in Palaeozoic spores, but is not precisely
defined. See also: capsula, cingulum, corona, patina, zona.
Foot layer (Fægri, 1956)
The inner layer of the ectexine. Synonym of
nexine 1, pedium and sole. See also: ectexine.
Foramen (pl. foramina, adj. forate, which is an abbreviation of foraminate) (Erdtman, 1952)
Synonym of pore. Comment: This term is sometimes used in its adjectival form
forate (e.g. Walker and Doyle, 1975) which is a synonym of pantoporate. A special form
of the term is internal foramen, which refers to features in the exine for example of
certain Compositae and Acanthaceae pollen grains. See also: internal foramen.
Fossaperturate (adj.) (Erdtman, 1952)

Refers to an equatorially aperturate, lobate pollen grain with the apertures
in the indentations between the lobes. Comment: The term belongs in the system of
ambs as described by Erdtman (1952). See also: angulaperturate, lobate,
sinu-aperturate.
Fossula (pl. fossulae, adj. fossulate) (Fægri and Iversen, 1950)

A feature of ornamentation consisting of an elongated, irregular groove in the
surface. See also: foveola.
Fossula (Kuprianova, 1948)

Synonym of areola.
Foveola (pl. foveolae, adj. foveolate) (Erdtman, 1952)

A feature of ornamentation consisting of more or less rounded depressions or
lumina more than 1 μm in diameter. The distance between foveolae is greater than their
breadth. See also: lumen, punctum.
Frustillum (pl. frustilla, adj. frustillate) (Fægri and Iversen, 1964)

Synonym of areola.
Furrow
A common word for an elongate aperture. Comment: Examples:
colpus, sulcus, tenuitas.
Furrow membrane (Wodehouse, 1935)
Synonym of colpus membrane.
Galea (pl. galeae, adj. galeate) (Sullivan, 1964)

A relatively large element of the outer wall of a spore consisting of a sharply
tapering spine and a broad bulbous base.
Garside's law/rule (Garside, 1946; Erdtman, 1952)

The name given to an unusual arrangement in developmental tetrads of
tri-aperturate pollen whereby the apertures form in groups of three at four points in the
tetrad. Comment: The name was given by Erdtman to a phenomenon described by Garside,
and apparently restricted to Proteaceae. See also: Fischer's law/rule.
Geminicolpate (adj.) (Erdtman, 1952)
Describing pollen grains with colpi arranged in pairs. See
also: pontoperculum.
Gemma (pl. gemmae, adj. gemmate) (Iversen and Troels-Smith, 1950)

A sexine element which is constricted at its base, higher than 1 μm, and that has
approximately the same width as its height.
Generative cell
The cell in a pollen grain which divides to form male gametes (Jackson, 1928).
Geniculum (pl. genicula, adj. geniculate) (Potonié, 1934)

A bulge in the equatorial exine of the colpus, often associated with a
separation of the sexine from the nexine and the rupturing of the latter. Examples:
Fraxinus (Oleaceae), Quercus (Fagaceae). Comment: If the separation forms a cavity the
term fastigium should be applied. See also: fastigium.
Germinal aperture (Wodehouse, 1935)

A hole in the furrow membrane through which the pollen tube emerges.
Comment: Examples: germ pore (synonym of pore), germinal furrow (synonym of
ectocolpus). Synonym of exitus.
Goniotreme (adj.) (Erdtman and Straka, 1961)
Synonym of angulaperturate.
Granular exine (Van Campo and Lugardon, 1973)

A type of exine stratification in which the infratectal layer is composed of more
or less rounded, granules rather than of columellae or other structures. See also:
columella, alveolate.
Granulate granulatus (adj.) (Erdtman, 1947)

With granules.
Granule (pl. granules, adj. granular, granulose)

General word for a small, rounded element.
Granulum (pl. granula, adj. granulate, granulose) (sensu Erdtman, 1952) A very small and
rounded element of the sexine/ectexine that is less than 1 μm in all directions. Comment: A
granulum is the diminutive for granum. See also: scabrate.
Granum (pl. grana) (Potonié, 1934)

The combined elements gemmae and verrucae.
Groove
A general descriptive word. See also: platea
luminosa, striate.
Gula (pl. gulae, adj. gulate) (Potonié and Kremp, 1955)

A rather ornate projecting, neck-like, extension on the proximal face of a trilete
spore. Example: Lagenicula horrida. Comment: This term is mainly used in the description
of some fossil megaspores.
Halo (Erdtman, 1952; Fægri and Iversen, 1989)

A clear zone around a well defined feature such as a spine or an aperture.
Examples: Ranunculus acris (Ranunculaceae), Valeriana (Valerianaceae).
Hamulate (adj.) (Krutzsch, 1959)

Describing a form of rugulate ornamentation consisting of irregularly arranged,
winding, or angular rounded muri of varying thickness, which do not form a distinct
reticulum, but rather a maze-like pattern. Example: Lycopodiella inundata
(Lycopodiaceae). Comment: Mostly used in spore terminology.
Harmomegathy (adj. harmomegathic) (Wodehouse, 1935)

The process by which pollen grains and spores change in shape to accommodate
variations in the volume of the cytoplasm caused by changing hydration.
H-endoaperture (Punt and Nienhuis, 1976)

An elaborate endoaperture, consisting of a central part which connects two
lateral, longitudinal elongations, forming an “H” shape. Examples: Cornus (Cornacaea),
Centaurium (Gentianaceae).
Heterobrochate (adj.) (Erdtman, 1952)
Describing a reticulum with brochi of different sizes. Examples: Adoxa
Moschatelina (Adoxaceae), Forsythia europaea (Oleaceae). See also: brochus, lumen.
Heterocolpate (adj.) (Iversen and Troels-Smith, 1950)

Describing pollen grains with both simple and compound colpi present.
Examples: Lythrum, Peplis (Lythraceae), Myosotis Boraginaceae). See also: pseudocolpus.
Heteropolar (adj.) (Erdtman, 1952)

Describing pollen or spores in which the distal and proximal faces of the exine
are different, either in shape, ornamentation or apertural system. Example: Echium vulgare
(Boraginaceae). Antonym: isopolar.
Heterosporous (adj.)
Describing plants producing both microspores and megaspores (Jackson,
1928). Antonym: homosporous. See also: isosporous.
Haploxylon-type (Rudolph, 1935)
Bisaccate pollen in which the outline of the sacci in polar view is more or less
continuous with the outline of the corpus, so that the grains appear a more or less smooth
ellipsoidal form. Examples: Pinus cembra, Picea (Pinaceae). See also: diploxylonoid,
haploxylonoid, Sylvestris-type.
Haploxylonoid (adj.) (Traverse, 1988)

Describing bisaccate pollen in which the outline of the sacci in polar view is
more or less continuous with the outline of the corpus, so that the grains appear a more or
less smooth ellipsoidal form. See also: diploxylonoid, Haploxylon-type,
Sylvestris-type.
Homosporous (adj.)
Describing plants producing only one kind of spores (Jackson, 1928).
Synonym of isosporous. Antonym: heterosporous.
Horn (Huynh, 1970)

An elongated part of an endoaperture, which is curved towards one of the poles.
Example: Anagallis arvensis (Primulaceae). See also: H-endoaperture.
Impression mark (Harley, 1998)

A mark on the proximal face of a pollen grain retained from the post-meiotic
stage. This mark can be linear from tetragonal tetrads or Y-shaped from tetrahedral tetrads.
Examples: Nypa fruticans, Howea balmooreana (Palmae).
In-A prefix used to emphasis the absence of a feature, as for example in inaperturate. See
also: A-.
Inaperturate (adj.) (Iversen and Troels-Smith, 1950)

Describing a pollen grain or spore without apertures. Example: Populus
(Salicaceae), Taxus (Taxaceae). Comment: The term should be used where apertures are
completely absent. If ectoapertures are absent, but endoapertures are present, the pollen is
cryptoaperturate. See also: cryptoaperture, omniaperturate.
Incidence (pl. incidences) (Batten and Christopher, 1981)

A structure formed by the termination of the inner wall layer at the endogerminal
which gives rise to a deep V-shaped atrium that points towards the centre of the grain.
Example: Pseudotrudopollis. Comment: This term is used in descriptions of the fossil
Normapolles group. See also: cuneus.
Hilum (pl. hila, adj. hilate) (Erdtman, 1952)
Circular, indistinctly delimited, irregular aperture or thinning in spores.
Examples: Aequitriradites verrucosus, Couperisporites tabulatus. Comment: This kind of
feature is present in certain bryophytes and fungi and may occur on the proximal or distal
face. See also: polumbra.
Homobrochate (adj.) (Erdtman, 1952)

Describing a reticulum with brochi of the same sizes. Example: Armeria
maritima (Plumbaginaceae). See also: brochus, lumen.
Inordinate (adj.) (Iversen and Troels-Smith, 1950)

Describing a pollen grain or spore with an arrangement of elements irregularly
distributed. Antonym: ordinate. Comment: The elements can be of structural or sculptural
origin. Examples: columellae under a tectum; scabrae on the tectum.
Insula (pl. insulae, adj. insulate) (Straka, 1964)

Synonym of areola.
Intectate (adj.) (Iversen and Troels-Smith, 1950)

Describing pollen grains without a tectum, but with sculpturing. Examples:
Viscum (Loranthaceae), Ilex (Araliaceae). See also: atectate, etectate, semitectate,
tectum.
Inter-
A prefix for in between. Comment: In palynology the prefix has been used in
many different contexts, from indicating position of apertures to defining areas of the
surface.
Interaspidium (pl. interaspidia) (Hoen and Punt, 1989)
The area of exine between aspides. Example: Dorstenia (Moraceae).
Infra-
A prefix meaning below or beneath. Comment: The term is often used for
patterns underneath a complete or partial tectum. Examples: infrareticulate,
infrarugulate, infrastriate, infrastructure, infratectate. See also: intra-.
Infratectum (pl. infratecta, adj. infratectate) (APLF, 1975)

A general term for the layer beneath the tectum, which may be alveolar,
granular, columellar, or structureless. Synonym of interstitium (Walker and Walker, 1981).
Infraturma (pl. infraturmae) (Potonié, 1956)

An artificial grouping of form-genera of spores and pollen in the
turma-system of Potonié. See also: anteturma, subturma, turma.
Inner tetrad mark (Gupta and Udar, 1986)

A tetrad mark at the inner layer of a spore and which does not reach up to the
margin when seen in polar view and always smaller than the outer tetrad mark.
Internal foramen (pl. internal foramina) (Skvarla and Larson, 1965)

Foramen present within the sexine/ectexine. See
also: foramen.
Internal tectum (Skvarla and Larson, 1965)

A more or less continuous layer within the outer sexine/ectexine composed of
laterally connected parts of columellae. Examples: Nigella damascena (Ranunculaceae),
Chrysantemum leucanthemum (Asteraceae). Comment: Differs from infratectum and
interstitium in being a single layer rather than referring to a number of elements under the
tectum.
Interporal lacuna (pl. interporal lacunae) (Wodehouse, 1935)

A lacuna occurring between the ends of two ectoapertures and adjacent to the
polar area of a lophate pollen grain. Example: Tragopogon pratensis (Compositae). See
also: lacuna.
Intercolpium (Fægri and Iversen, 1950)
The meridional segment between two colpi and extending to the poles.
Comment: The terms mesocolpium and apocolpium are recommended because they
enable a more precise subdivision of the pollen surface. See also: apocolpium,
mesocolpium.
Interlacunar gaps (Wodehouse, 1935)

Gaps in the short sexinous ridges which often divide the ectoapertures of lophate
pollen grains into lacunae. Example: Cichorium intybus (Compositae). See also: lophate.
Interlacunar ridges (Wodehouse, 1935)

Ridges separating the lacunae in lophate pollen grains. Example:
Cichorium intybus (Compositae).
Interloculum (pl. interlocula) (Thomson and Pflug, 1953)

A space between the inner- and outer wall layers that is present around most or
all of the pollen grain. Example: Extratriporopollenites conjunctus. Comment: The term
was originally used for features in pollen of the Triatrioaperturate category of the fossil
Normapolles group.
Intra-A prefix meaning within. See also:
infra-.
Intrareticulum (adj. intrareticulate) (Iversen and Troels-Smith, 1950)

Synonym of infrareticulum.
Isodiametric tetrad
A general term for tetrads in which all members are more or less the same in
size. Antonym: anisodiametric tetrad.
Isopolar (adj.) (Erdtman, 1947)

Describing a pollen grain or spore in which the proximal and distal faces of the
exine are alike. Antonym: heteropolar.
Interporium (pl. interporia) (Iversen and Troels-Smith, 1950)
The meridional segment between two pores extending to the poles.
Comment: The terms mesoporium and apoporium are recommended because they
enable a more precise subdivision of the pollen surface. See also: apoporium,
mesoporium.
Interradial (adj.) (Couper and Grebe, 1961)

Referring to areas of the proximal face or the equatorial periphery of trilete
spores, lying between the radial arms of the laesurae. Comment: Interradial is also used to
indicate the position of apertures with reference to tetrad organisation (Walker and Doyle,
1975).
Interstitium (Walker and Walker, 1981)

The layer of the exine situated between the nexine and the tectum.
Synonym of infratectum.
Intexine (Potonié, 1934)

Synonym of nexine.
Intine (Fritzsche, 1837)

The innermost of the major layers of the pollen grain wall underlying the exine
and bordering the surface of the cytoplasm. Comment: The intine is not acetolysis resistant
and is therefore absent in conventionally prepared palynological material.
Laesura (pl. laesurae, suffix-lete) (Erdtman, 1946) The arm of a proximal fissura or scar
of a spore. Comment: A monolete spore has one laesura, a trilete spore three (although
some palynologists consider a trilete spore to have a single triradiate laesura). A laesura
comprises a commissure which may be bordered by a margo (labrum). See also:
monolete, trilete.
Isospore
A spore of a plant producing only one kind of spore (Jackson, 1928). See also:
miospore.
Isosporous (adj.)
Describing plants producing only one kind of spore (Jackson, 1928). Synonym
of homosporous. Antonym: heterosporous.
Kyrtome (Potonié and Kremp, 1955)

A more or less arcuate fold or band in the interradial areas outside the laesurae
of trilete spores. Examples: Ahrensisporites guerickei, Concavisporites rugulatus.
Comment: Some palynologists prefer to use torus for separate interradial bands, and
kyrtome for a connected feature. See also: torus.
Labrum (pl. labra, adj. labrate) (Thomson and Pflug, 1953)

Protruding part of the exine at the pores of porate pollen grains. Example:
Lusatipollis. Comment: This feature is especially prominent in many representatives of the
fossil pollen of the Normapolles group. See also: vestibulum.
Labrum (pl. labra, adj. labrate) (Couper and Grebe, 1961)

The elevated and/or thickened part of a laesura between the commissure and
the remainder of the proximal surface. See also: margo. Comment: The term is mainly
used in the descriptions of fossil spores. Because of possible confusion with the term
labrum (sensu Thomson and Pflug, 1953) it is, however, suggested to use the term margo
instead.
Lacuna (pl. lacunae, adj. lacunate) (Wodehouse, 1928)

A depressed area surrounded by ridges in lophate pollen grains. See also:
abporal lacuna, circumpolar lacuna, equatorial lacuna, interporal lacuna,
paraporal lacuna, polar lacuna, poral lacuna.
-lete (Erdtman, 1969)
A suffix to denote the presence (or absence) of laesura. Examples: alete,
monolete, trilete.
Laevigate (adj.)
A general term for smooth, as if polished (Jackson, 1928). Synonym of
psilate. Orthographical variant: levigate. Comment: Although the correct Latin spelling
is levigate, the spelling laevigate is accepted throughout the botanical literature (Jackson,
1928). The term has mostly been used in the descriptions of fossil spores. Example:
Laevigatisporites.
Lalongate (adj.) (Erdtman, 1952)

Describing the shape of a transversely elongated endoaperture. Example:
Filipendula (Rosaceae). See also: lolongate.
Lamella (pl. lamellae, adj. lamellar, lamellate)

A general term for a thin layer (Jackson, 1928).
Latimurate (adj.) (Erdtman, 1952)

Describing a pollen grain or spore with broad muri.
Latiporate (adj.) (Norem, 1958)

Describing pollen grains with pores in one hemisphere only. Examples: Juglans,
Carya (Juglandaceae).
Latitudinal (adj.)
A general descriptive term, in palynology applied to features which run in lines
parallel to the equator. Antonym: longitudinal. See also: sulcus.
Layer
A general term. Applied in palynology to any distinct stratum of the
sporoderm (APLF, 1975).
-lept (Erdtman, 1969)

A suffix for thin, indicating that a pollen grain has a leptoma.
Leptoma (Erdtman and Straka, 1961)

A thin area at the distal pole of a pollen grain, presumed to function as an
aperture. Example: Pinus (Pinaceae). Comment: substitute term for cappula. See also:
tenuitas.
Levigate (adj.)
Orthographical variant of laevigate.
Limb (Wodehouse, 1935)

Synonym of equatorial outline.
Limbus (Potonié and Kremp, 1955)
A sharp narrow crease of the saccus or pseudosaccus where the outer and inner
exine layers are fused. Example: Nuskoisporites dulhuntyi.
Linear tetrad (Erdtman, 1945b)

A uniplanar tetrad in which the four members are arranged in a row. Example:
Typha spp. (Typhaceae). See also: tetrad.
Lira (pl. lirae) (Erdtman, 1952)

A narrow ridge which forms the murus in a striate pattern. Synonym
of murus and vallum.
LO-analysis (Erdtman, 1952)

A method for analyzing patterns of sexine organization by means of light
microscopy. Comment: This method is valuable for elucidating exine patterns. When
focused at high level (H), raised sexine elements appear bright (Lux), whereas holes in the
tectum are relatively dark (Obscuritas). At lower focus (L) holes become lighter and the
sexine elements become darker. See also: LO-pattern, OL-pattern.
Lobate (adj.) (Kuyl et al., 1955)

Describing an equatorially aperturate pollen grain with a lobed shape in polar
view. Comment: Belongs to a system of shapes combined with the position of the apertures
as introduced by Kuyl et al.
Lolongate (adj.) (Erdtman, 1952)

Describing the shape of a longitudinally elongated endoaperture. Example:
Rumex spp. (Polygonaceae). See also: lalongate.
Longi-
A prefix for long.
Longiaxe (adj. logiaxal) (Van Campo, 1966)

Pollen grains with a polar axis longer than their equatorial diameter. Synonym of
prolate (s.l.). Antonym: breviaxe. Comment: Thompson and Pflug (1953) recognised
Longaxones as a group of mid-Cretaceous and younger angiosperm pollen contrasted with
Brevaxones.
Longitudinal (adj.)
A general descriptive term, in palynology applied to features which run in lines
between the poles. Antonym: latitudinal. See also: lolongate.
LO-pattern (Erdtman, 1952)

A pattern of ornamentation that appears to show “bright islands” at high focus (H)
and that become dark at low focus (L), observed when using LO-analysis. Comment: The
reverse of OL-pattern.
Lophate (adj.) (Wodehouse, 1935)

Describing a pollen grain in which the outer exine is raised in a pattern of ridges
(lophae) surrounding depressions (lacunae). Example: Hieracium (Compositae). See also:
echinolophate, psilolophate.
Loxocolp(or)ate (adj.) (Erdtman and Straka, 1961)

Describing zonocolp(or)ate pollen with ectocolpi arranged so that they converge in
pairs. Comment: Such apertures generally occur in aberrant grains with more than the typical
number of apertures.
Lumen (pl. lumina) (Potonié, 1934)

The space enclosed by the muri. See
also: brochus, reticulum.
Macrospore
General term for the larger spores of heterosporous vascular plants (Jackson,
1928). Antonyms: microspore (in the general botanical sense); miospore (in the sense
of Guennel, 1952). Comment: In dispersed fossil spores where it is not always possible to
determine whether the parent plants were heterosporous the term indicates size only
(usually >200 μm). See also: megaspore.
Maculate (adj.) (Potonié, 1934)
A general descriptive term. In palynology it can be applied to pollen grains or
spores with a spotted exine, often due to variations in internal exine structure. Example:
Maculatisporites.
Margo (pl. margines, adj. marginate) (Iversen and Troels-Smith, 1950) An area of exine
around an ectocolpus that is differentiated from the remainder of the sexine, either in
ornamentation or by difference in thickness. See also: annulus, labrum.
Massa (pl. massae) (sensu Potonié, 1956)

A specialised structure composed of aborted spores and tapetal material found
on certain megaspores and often referred to as a floating apparatus. Applied only to
megaspores. Examples: Azolla (Azollaceae), Cytosporites varius.
Massula (pl. massulae, adj. massulate)

A general term for aggregations of pollen grains dispersed as a unit (Jackson,
1928). Examples: Mimosaceae, Periplocaceae. See also: dispersal unit, polyad,
pollinium.
Medine (Saad, 1963)

A term applied to a faintly lamellated, acetolysis resistant layer
considered to be situated between intine and exine. See also: mesine.
Megaspore
A general term for large spores of heterosporous vascular plants (Jackson,
1928). See also: microspore, macrospore.
Meiosis (adj. meiotic)

The reduction division of chromosomes (Jackson, 1928).
Meiospore
General term for a spore produced by meiosis (Jackson, 1928).
Melissopalynology (Maurizio and Louveaux, 1960)

The study of pollen grains collected by bees, and/or found in honey. Comment:
The variants melitopalynology (Erdtman, 1952) and melittopalynology (Fægri and Iversen,
1975) are sometimes encountered.
Membrana colpi (pl. membranae colpi) (Iversen and Troels-Smith, 1950)

Synonym of colpus membrane.
Membrana pori (pl. membranae pori) (Iversen and Troels-Smith, 1950)
Synonym of pore membrane.
Meridional (adj.) (Fægri and Iversen, 1950)

Describing longitudinal features on the surface of a pollen grain or spore which
run along lines perpendicular to the equator.
Meridionosulcus (adj. meridionosulcate) (Sampson, 1976)

A meridional sulcus. Example: Degeneria vitiensis (Degeneriaceae). See
also: zonasulcus.
Mesine (Rowley, 1959)

A laminated, electron-dense layer considered to lie between the intine and the
exine. Comment: Probably the same as the medine, but defined on different methods of
microscopy. See also: medine.
Meso-
A prefix meaning middle.
Mesoaperture (adj. mesoaperturate) (Thanikaimoni, 1980)

The middle part of a compound aperture in which there is also an
ectoaperture and an endoaperture. Example: Polygonum aviculare (Polygonaceae).
Mesocolpium (pl. mesocolpia) and Mesoporium (pl. mesoporia) (Erdtman,

1952)
The area of a pollen grain surface delimited by lines between the apices of
adjacent colpi or the margins of adjacent pores. See also: apocolpium, apoporium,
intercolpium, interporium.
Micro-
A prefix for small. Comment: In palynology, generally used to denote features
less than 1 μm. Examples: microechinate, microverruca. See also: nano-.
Microreticulum (adj. microreticulate) (Praglowski and Punt, 1973)

A reticulate ornamentation consisting of muri and lumina smaller than 1 μm. See
also: reticulum.
Microspore
A general term for the smaller spores of heterosporous plants, that is, the spores
from which the microgametophyte develops (Jackson, 1928). Antonyms: macrospore,
megaspore. Comment: The term is sometimes used in palaeopalynology for small spores
generally, whether homospores, microspores or even small megaspores. This usage should
be avoided, in favour of the term miospore or small spore, but only if it is not known the
plant is homosporous or not. A pollen grain is the microgametophyte of a seed plant. In
developmental studies, a pollen grain is sometimes referred to as microspore until
microspore mitosis. See also: miospore, pollen.
Microsporocyte
A general term for the mother cell of a microspore or pollen grain (Jackson,
1928). See also: pollen mother cell.
Middle part (Punt and Rentrop, 1973)

Synonym of mesoaperture.
Metareticulum (adj. microreticulate) (Borsch and Bathlott, 1998)

A reticulum which is characterized by the consistent presence of one porate
aperture in each lumen. Examples: Froelichia floridana (Amaranthaceae), Viviania rosea
(Vivianiaceae), Kallstroemia maxima (Zygophyllaceae).
Miospore (Guennel, 1952)
A general term for all fossil plant spores smaller than 200 μm, regardless of
whether they are isospores, microspores, small megaspores, prepollen or pollen grains.
Antonym: macrospore. See also: microspore, pollen.
Mitosis
A general term for nuclear division involving no reduction of chromosomes
(Jackson, 1928).
Monolete (adj.) (Erdtman, 1943)
Describing a spore with a single laesura. Example: Dryopteris
(Dryopteridaceae). See also: trilete.
Monosaccate (adj.) (Potonié and Kremp, 1954)

Describing a pollen grain with a single saccus. Example: Florinites
antiquus. See also: bisaccate.
Morphon (Van der Zwan, 1979)

A group of form-species exhibiting continuous variation of morpholo gical
characteristics in a single time sequence (“horizontal” variation). Comment: Many authors
use the word “complex” in a very similar way. See also: palynodeme.
Multibaculate (adj.) (Erdtman, 1952) Synonym of

pluricolumellate.
Multiplanar tetrad (Walker and Doyle, 1975)

A tetrad in which the individual members are arranged in more than one plane.
Comment: Decussate and tetrahedral tetrads are multiplanar, whereas rhomboidal, linear
and T-shaped tetrads are uniplanar.
Murornate sculpture elements (Smith and Butterworth, 1967)

Elevations of the general surface. Examples: cristae, muri.
Monad (Selling, 1947)
A pollen grain or spore dispersed as an individual unit, rather than in
association with others, such as in a dyad, tetrad or polyad.
Mono-
A prefix for one.
Monoaperturate (adj.)
Describing a pollen grain or spore with a single aperture. Comment:
Examples: monocolpate (Iversen and Troels-Smith, 1950), monoporate (Iversen
and Troels-Smith, 1950) monosulcate (Erdtman, 1952).
Nexine 1 (Erdtman, 1966a)

Synonym of foot layer, pedium, sole.
Nexine 2 (Erdtman, 1966a) Synonym of

endexine.
Murus (pl. muri) (Erdtman, 1943)

A ridge that is part of the ornamentation and, for example, separates the lumina
in a reticulate pollen grain or the striae in striate pollen grain. Comment: Muri in striate
patterns are sometimes called valla (Iversen and Troels-Smith, 1950) or lirae (Erdtman,
1952).
Nano- (Erdtman, 1969)

A prefix for elements smaller than 0.5 μm. See also:
micro-.
Negative reticulum (Erdtman, 1943)

A general term used to describe patterns of ornamentation in which sexine
areas are separated by narrow, reticulately arranged grooves. See also: areola,
frustillum.
Nexine (Erdtman, 1952)

The inner, non-sculptured part of the exine which lies below the sexine.
Antonym: sexine. Comment: Although the term endexine as originally defined (sensu
Erdtman, 1943) was more or less synonymous with nexine it no longer is because endexine
is now invariably used in the sense of Fægri (1956). Nexine and sexine are distinguished on
purely morphological criteria, whereas ectexine and endexine differ in their staining
properties. The two sets of terms therefore have slightly different applications. See also:
endexine (sensu Fægri, 1956).
Non-aperturate (adj.) (Fægri and Iversen, 1950)

Synonym of inaperturate.
Normapolles (Pflug, 1953)

A group of Cretaceous and Lower Palaeogene pollen, usually triporate, with a
complex pore apparatus.
NPC-classification (Erdtman and Straka, 1961)

A morphological system for classifying pollen grains and spores that was based
on the number, position and character of their apertures.
Nudate (adj.) (Punt et al., 1976)

Synonym of psilate.
Oculus (pl. oculi, adj. oculate) (Thomson and Pflug, 1953)
The much enlarged part of the pore structure in pollen of the fossil Normapolles
group. Example: Oculopollis. Comment: The enlarged part is a swelling of the outer wall
layer on one or both surfaces of a grain in the region of an exogerminal. An annulus is
uniformly thick and completely surrounds the exogerminal region. See also: annulus.
OL-pattern (Erdtman, 1952)

A pattern of ornamentation that appears to show “dark islands” at high focus
(H) and that become bright at low focus (L). Comment: The reverse of a LO-pattern.
Omniaperturate (adj.) (Thanikaimoni, 1984)

Describing a pollen grain in which the exine is very thin or absent and the intine
is thick, so that no specific apertural region can be distinguished and thus the whole surface
can be considered apertural in nature.
Oncus (pl. onci) (Hyde, 1955)

A lens-shaped structure that is not resistant to acetolysis and occurs beneath
the apertures of many kinds of pollen grains. Example: Corylus (Betulaceae). See also:
Zwischenkörper.
Oblate (adj.) (Erdtman, 1943)
Describing the shape of a pollen grain or spore in which the polar axis is shorter
than the equatorial diameter. Comment: This term belongs to the system of shape classes
suggested by Erdtman (1943, extended in 1952), based on the measurements of the polar
axis (P) and equatorial diameter (E). In this system oblate is defined as a ratio between the
polar axis and the equatorial diameter of 0.50–0.75. See also: oblate spheroidal, P/E
ratio, peroblate, perprolate, prolate, prolate spheroidal, shape classes,
spheroidal, suboblate, subprolate, subspheroidal.
Oblate spheroidal (adj.) (Erdtman, 1952)

Describing the shape of a pollen grain or spore in which the ratio between
the polar axis and the equatorial diameter is 0.88–1.00. See also: oblate, P/E ratio,
peroblate, perprolate, prolate, prolate spheroidal, shape classes,
Ocellus (pl. ocelli, adj. ocellate) (Grebe, 1971)

Synonym of cata-ulcus. See
also: ulcus.
Optical (cross-) section

The image seen in optical microscopy when the plane of focus is half way
through a palynomorph.
Orbicule (pl. orbicules, adj. orbicular)

A general term, applied in palynology for an orbicular granule of
sporopollenin. Synonym of Ubisch body.
Ordinate (adj.) (Iversen and Troels-Smith, 1950)

Describing a pollen grain or spore with an arrangement of elements regularly
distributed. Antonym: inordinate. Comment: The elements can be of structural or
sculptural origin. Examples: columellae under a tectum forming a reticulum (example:
Vaccaria pyramidata, Caryophyllaceae); scabrae on the tectum (example: Alchemilla
glabra, Rosaceae) arranged in a regular pattern.
Ornamentation (Potonié, 1934)

A general term that is useful for describing the organisation of features. See also:
pattern, sculpture.
Ornate (adj.) (Erdtman, 1953)

Describing a reticulate ornamentation consisting of broad, curved muri and
lumina that are often anastomosing. Example: Ceiba aesculifolia (Bombacaceae).
Orthocolpate (adj.) (Erdtman and Straka, 1961)
Describing a pollen grain with the colpi in the most common position,
perpendicular to the equator. Comment: This term is not needed in most instances, where
zonocolpate would suffice, but exists to contrast with the term loxocolpate.
Os (pl. ora, adj. orate) (Erdtman, 1952)

Synonym of endoaperture. Comment: The term is now mainly used in so far
as that ora is a component of the terms colporus and pororate.
Operculum (pl. opercula, adj. operculate) (Wodehouse, 1935)
A distinctly delimited sexineectexine structure which covers part of an
ectoaperture and which is completely isolated from the rest of the sexine. See also:
pontoperculum.
Palaeopalynology (Manten, 1966)

The study of fossil palynomorphs.
Antonym: actuopalynology.
Palynodebris (Manum, 1976)

All palynomorph-sized particles in a sediment excluding those that actually
are palynomorphs but including, for example, wood fragments, cuticles and some animal
remains. See also: phytoclast.
Palynodeme (Visscher, 1971)

A group of palynomorph species (form-species) that intergrade and represent
the reflection of a known or hypothetical plant species. Comment: As originally used the
concept was phylogenetic and referred to characters changing in time (“vertical” change).
The term is, however, misused by some authors as if synonymous with morphon and the
less formal term “complex”. See also: morphon.
Palynofacies (Combaz, 1964)

The assemblage of phytoclasts found in a particular sediment, such as
palynomorphs, wood fragments, cuticles, etc. Comment: The term is actually used in two
senses, namely the palynolithofacies and palynobiofacies. See also: phytoclast.
Palynogram (Erdtman, 1952)

A diagram summarising the main morphological features of a palynomorph.
Palynology (Hyde and Williams, 1944)

The study of pollen grains and spores and of other biological materials that can
be studied by means of palynological techniques. Comment: A number of subdisciplines
may be recognised, including palaeopalynology, aeropalynology,
melissopalynology and pollen analysis.
Outline
A general descriptive word. Applied in descriptive terms like equatorial
outline and outline in polar view.
P/E ratio (Erdtman, 1943)

The ratio of the length of the polar axis (P) to the equatorial diameter
(E). Comment: Erdtman suggested a widely used system of shape classes defined on the
basis of P/E ratios. See also: oblate, oblate spheroidal, peroblate, perprolate,
prolate, prolate spheroidal, shape classes, spheroidal, suboblate,
subprolate, subspheroidal.
Papilla (pl. papillae, adj. papillate) (Traverse, 1955)
A general term, applied in palynology to parallel sided exinous elements with
rounded apices, less than 1 μm in length. See also: scabrate.
Paracavate (adj.) (Balme, 1988)

An exine in which the intexine is clearly defined but in which its degree of
separation from the exoexine is uncertain or indeterminate. Example: Ancyrospora langii.
Paraisopolar (Praglowski et al., 1983)

Describing a pollen grain whose polar faces differ only in the attachment of
viscin threads to the proximal pole. Synonym of subisopolar. Comment: Most pollen
grains with viscin threads have polar faces of which one is less/more convex than the other.
Paraporal lacuna (pl. paraporal lacunae) (Wodehouse, 1935)

A lacuna of a lophate pollen grain lying in the mesocolpial region adjacent to one
side of an equatorial ridge. Example: Taraxacum officinale (Compositae). See also:
lacuna.
Palynomorph (Tschudy, 1961)
A general term for all entities found in palynological preparations. Comment: In
addition to pollen grains and spores, the term encompasses acritarchs, dinoflagellates and
scolecodonts, but not other microfossils, such as diatoms, that are dissolved by
hydrofluoric acid.
Panporate (adj.) (Erdtman and Vishnu-Mittre, 1956)

Synonym of pantoporate.
Panto-, Pan-
A prefix for global distribution.
Synonym of peri-.
Pantoaperturate (adj.) (Erdtman and Vishnu-Mittre, 1956)

Describing a pollen grain with apertures spread over the surface sometimes
forming a regular pattern. Comment: Such pollen grains may be, for example,
pantocolpate, pantocolporate or pantoporate. The terms based on the prefix peri-
as used in the classification of pollen types of Iversen and Troels-Smith (1950) are not
recommended.
Papilla (pl. papillae, adj. papillate) (Wodehouse, 1935)

A small protuberance. Comment: The term is mostly used in describing pollen
of Taxodiaceae (Gymnospermae).
Per-A prefix for 1. Extremely, and 2. Through. Comment: In palynological terms
the prefix is often used for complete (e.g. in pertectate) or very (e.g. in peroblate).
Pererect (adj.) (Reitsma, 1970) Synonym of

perprolate.
Paraporal ridge (Wodehouse, 1935)
A ridge bounding a pore, extending in a meridional direction. Applied to a
lophate pollen grain. Examples: Tragopogon pratensis, Taraxacum officinale
(Composiatae).
Parasyncolpate (adj.) (Erdtman, 1952) and parasyncolporate (Van der Ham, 1977)
Describing syncolp(or)ate pollen grains in which the apices of the colpi
divide into two branches and anastomose towards the poles, delimiting an isolated area
known as the apocolpial field. Examples: Nymphoides peltata (Menyanthaceae), Eugenia
(Myrtaceae). See also: syncolp(or)ate, apocolpial field.
Patella (pl. patellae, adj. patellate) (Pocock, 1961a)

Synonym of patina. See also: capsula,
cingulum.
Patina (adj. patinate) (Butterworth and Williams, 1958)

A thickening of the exine of spores that extends over the entire surface of one
hemisphere. Example: Cingulatisporites, Patellasporites.
Pattern
A general word, applied in palynology either to surface features or
infratectal elements, such as columellae.
Pedium (Erdtman, 1966b) Synonym of

foot layer.
Perforate (adj.) (Iversen and Troels-Smith, 1950)
A general adjective indicating the presence of holes, applied in palynology to
holes less than 1 μm in diameter and generally situated in the tectum. See also: punctum,
scrobiculus.
Peri-Synonym of panto-. Comment: Iversen and Troels-Smith (1950) used the term peri-
(as for example, in pericolpate, periporate and pericolporate) in their classification of
pollen types, but terms based on panto- are much more widely used. See also:
pantoaperturate.
Perine (Erdtman, 1943)

A sporoderm layer that is not always acetolysis resistant and is situated around
the exine of many spores. Example: Pteridium (Hypolepidaceae). Synonym of perispore.
Comment: The term perine should be used in conjunction with sexine and nexine, whereas
perispore should be used with exospore and endospore.
Perinium (Jackson, 1928) Synonym of perine. Comment: A

latinised form of the term perine.
Perispore (Russow, 1872)

Synonym of perine. See also:
endospore, exospore.
Perisporium (Erdtman, 1943)

Synonym of perine. Comment: A latinised form of the
term perispore.
Peritreme (adj.) (Erdtman and Straka, 1961)

Describing a pollen grain with equatorial apertures situated around an outline
that is circular in polar view. Comment: The term is not recommended because it is based
upon the suffix -treme.
Peroblate (adj.) (Erdtman, 1943)

Describing the shape of a pollen grain or spore in which the ratio between the
polar axis and the equatorial diameter is less than 0.50. See also: oblate, oblate
spheroidal, P/E ratio, perprolate, prolate, prolate spheroidal, shape classes,
Perprolate (adj.) (Erdtman, 1943)
Describing the shape of a pollen grain or spore in which the ratio between the polar
axis and the equatorial diameter is more than 2. See also: oblate, oblate spheroidal, P/E
ratio, peroblate, prolate, prolate spheroidal, shape classes, spheroidal,
suboblate, subprolate, subspheroidal.
Per-reticulate (adj.) (Fægri and Iversen, 1975)

Structural elements fused distally forming an open reticulum. Comment: It applies
to the same structure as eureticulate, but is defined on a different basis.
Pertectate (adj.) (Erdtman, 1969)

Synonym of eutectate.
Pertransverse (adj.) (Reitsma, 1970)
Synonym of perprolate.
Phytoclast (pl. phytoclasts)

A general term for plant-derived, more or less resistant-walled, particle
occurring in a sediment. See also: palynodebris.
Pilum (pl. pila, adj. pilate) (Erdtman, 1952)

A sexine element, usually standing directly on the nexine, consisting of a rod-like
part (columella) and a swollen apical part (caput).
Pitted (adj.)
A general term for small depressions (Jackson, 1928).
Synonym of foveolate (in palynology).
Planaperturate (adj.) (Erdtman, 1952)

Describing a pollen grain with an angular outline, in which the apertures are
situated in the middle of the sides when seen in polar view, rather than at the angles. Example:
Tilia (Tiliaceae). Antonym: angulaperturate. Comment: The term is useful for describing
the position of apertures, but should be avoided as a description of equatorial outline.
Platea luminosa (pl. plateae luminosae) (Iversen and Troels-Smith, 1950)

Synonym of groove. Comment: The term was introduced for use in striate
pollen and to contrast with lumina, which was restricted to reticulate pollen.
Pleurotreme (adj.) (Erdtman and Straka, 1961)

Synonym of planaperturate.
Plica (pl. plicae, adj. plicate) (Thomson and Pflug, 1953)
A general term for a fold, applied in palynology to ridge-like folds of the exine in
Ephedra (Ephedraceae) and Lusatipollis. See also: polyplicate, taenia.
Pluricolumellate (adj.) (Reitsma, 1970)

With the columellae arranged in several rows beneath each murus. See also:
duplicolumellate, simplicolumellate.
Polar area (Iversen and Troels-Smith, 1950)

Synonym of apocolpium. Comment: In spores often used for
the area around a pole.
Polar area index (PAI) (pl. polar area indices) (Iversen and Troels-Smith, 1950)
Synonym of apocolpium index.
Platea (pl. plateae) (Thomson and Pflug, 1953)
Areas of the inner wall layer separated by a triradiate channel extending between
the endogerminals of a Normapolles pollen grain. Example: Pompeckjoidaepollenites.
Polar view (Erdtman, 1943)
A view of a pollen grain or spore in which the polar axis is directed towards the
observer. Antonym: equatorial view. See also: amb.
Polarity
The condition of having distinct poles (Jackson, 1928). Comment: The polarity
of palynomorphs may be determined from their orientation in tetrads, or by inference from
the distribution of apertures, or other features. See also: apolar, heteropolar, isopolar.
Pole (Wodehouse, 1935)

Either of the two extremities of the polar axis. See
also: polar axis.
Polar axis (pl. polar axes) (Wodehouse, 1935)
The straight line between the distal and proximal poles of a pollen grain or
spore. See also: equatorial diameter, polarity.
Polar distance (Punt, 1984)

The vertical distance from the equator to the pole (pd).
Polar field (Beug, 1961)

Synonym of apocolpium.
Polar lacuna (pl. polar lacunae) (Wodehouse, 1928)

A lacuna situated at the pole of a lophate pollen grain. See
also: lacuna, lophate.
Pollen (Linnaeus, 1750)
The microgametophyte of seed plants, developed from the microspore. See also:
spore, microspore.
Pollen analysis (Von Post, 1916)

The study of assemblages of dispersed palynomorphs such as those isolated
from samples of peat. See also: palynology.
Pollen cement (Heslop-Harrison, 1968)

Synonym of pollenkitt.
Pollen class (pl. pollen classes) (Fægri and Iversen, 1950)

An artificial grouping of pollen grains that share a distinctive character, or suite
of characters. Such classes are useful in identification keys and may be subdivided into
more restrictive categories, pollen types and pollen groups. Example: tricolpate class.
Pollen coat (Knox, 1984)

Synonym of pollenkitt.
Pollen group (Punt, 1971)

A pollen morphological category, subsidiary to a pollen type, including a
number of pollen grains that show intergrading characters but no distinguishing characters.
See also: pollen class, pollen type.
Pollenkitt (Knoll, 1930)

A sticky material, produced by the tapetum, that may hold pollen grains together
during dispersal. See also: tryphine.
Pollen mother cell

Synonym of microsporocyte.
Pollen type (Punt, 1971)

A pollen morphological category, subsidiary to a pollen class, and including
pollen grains which can be distinguished either by one distinct character or by a unique
combination of characters. See also: pollen class, pollen group.
Pollinium (pl. pollinia)

A general term for aggregations of many pollen grains, which form dispersal
units (Jackson, 1928). Examples: Asclepiadaceae, Orchidaceae. See also: dispersal unit,
massula.
Polumbra (Balme, 1988)

A darkened triangular or subcircular area centred on the proximal pole. Example:
Retusotriletes distinctus. Comment: The feature appears to be most commonly observed in
specimens that have lost a perisporal outerexoexinal layer. See also: hilum.
Poral lacuna (pl. poral lacunae) (Wodehouse, 1928)
A lacuna of a lophate pollen grain surrounding an endoaperture, which
communicates with adjacent abporal lacunae via interlacunar gaps. Example: Cichorium
intybus (Compositae). See also: lacuna, lophate.
Polyad (Iversen and Troels-Smith, 1950)
A dispersal unit comprising more than four pollen grains. Example: Acacia
(Mimosaceae).
Polyannulus (pl. polyannuli, adj. polyannulate) (Batten and Christopher, 1981)
A structure in which the sexine of the outer aperture has multiple layers each
with its own thickening. Example: Atlantopollis. Comment: This term refers especially to
pollen grains of the fossil Normapolles group.
Polychotomosulcate (adj.) (Walker and Walker, 1986)

Describing a pollen grain with a many branched sulcus.
Polyplicate (adj.) (Iversen and Troels-Smith, 1950)

Describing a pollen grain with more than three meridional ridges (plicae)
separated by deep grooves. Example: Ephedra (Ephedraceae). See also: plica, striate,
taenia.
Polyporate (adj.) (Moar, 1993)

With many pores. Synonym of pantoporate
and zonoporate.
Pontoperculum (pl. pontopercula, adj. pontoperculate) (Erdtman, 1952)

A type of operculum that is not completely isolated from the remainder of the
sexine but linked to it at the ends of the aperture. Example: Sanguisorba officinalis
(Rosaceae), Passieflora tetandra (Passiefloraceae). See also: geminicolpate.
Pore (pl. pores, adj. porate) (Jackson, 1928; Wodehouse, 1935)
A general term, applied in palynology to a circular or elliptic aperture with a
length/breadth ratio less than 2. See also: aperture, porus, ulcus.
Pore canal (Thomson and Pflug, 1953)

The space between the ectopore and the endopore. See also:
aspis channel.
Pore membrane (Wodehouse, 1935)
The aperture membrane of a pore.
Poro-colpate (adj.) (Clarke et al., 1979)

Describing a pollen grain with an arrangement of apertures in which colpi
alternate with pores round the equator. Example: Pardoglossum (Boraginaceae).
Pororate (adj.) (Erdtman, 1952)

Describing a pollen grain with compound apertures in which both the
ectoaperture and the endoaperture are pores and the two are not congruent. Example:
Myrica gale (Myricaceae). Comment: Pollen grains that have congruent ectopores and
endopores are generally simply referred to as porate. See also: colporate.
Porus (pl. pori, adj. porate) (Potonié, 1934)

Synonym of pore.
Pre-A prefix for before. Comment: As for example in prepollen,
pretectum.
Prepollen (Renault, 1896; Chaloner, 1970)

The microspores of certain extinct seed plants characterised by proximal
apertures and presumed proximal germination, rather than the distal, equatorial or other
typical apertures of seed plant pollen grains.
Primexine (Heslop-Harrison, 1963)
A developmental precursor of at least part of the exine (the sexine/
ectexine), formed during the tetrad stage that is composed largely of polysaccharides
and therefore lacks resistance to acetolysis.
Pro- (Heslop-Harrison, 1963)

A prefix indicating a developmental precursor of a feature. Example:
probaculum.
Projectate (adj.) (Mtchedlishvilli, 1961)

Describing a pollen grain in which the apertures are borne on the ends of
strongly projecting arms. Example: Aquilapollenites. See also: triprojectate.
Prolate (Erdtman, 1943)

Describing the shape of a pollen grain or spore in which the polar axis is larger
than the equatorial diameter. Comment: This term belongs to the system of shape classes
suggested by Erdtman (1943, and extended in 1952), based on the measurements of the
polar axis (P) and equatorial diameter (E). In this system prolate is defined as a ratio
between the polar axis and the equatorial diameter of 1.33– 2.00. See also: oblate,
oblate spheroidal, P/E ratio, peroblate, perprolate, prolate sp heroidal, shape
classes, spheroidal, suboblate, subprolate, subspheroidal.
Prolate spheroidal (Erdtman, 1952)

polar axis and the equatorial diameter is 1.00–1.14. See also: oblate, oblate
spheroidal, P/E ratio, peroblate, perprolate, prolate, shape classes,
Protosaccus (pl. protosacci, adj. protosaccate) (Scheuring, 1974)

A saccus which is completely filled with an alveolar structure. This feature is
used in the description of pollen in the Perm-Trias. Example: Lueckisporites virkkiae.
Comment: For extant saccate pollen grains don't show this character it is considered to
be primitive. See also: bisaccate, monosaccate, pseudosaccus, saccus.
Proximal
A common descriptive term (Jackson, 1928) used in contrast to distal, applied
in palynology to features on the surface that faces towards the centre of the tetrad during
development (Wodehouse, 1935). Antonym: distal. See also: ana-, cata-, polarity.
Proximal pole (Wodehouse, 1935) The centre of
the proximal face. Antonym: distal pole. See also:
proximal.
Proximocavate (adj.) (Balme, 1988)

An exine in which the exoexine is detached, or partly detached, from the intexine
only on the proximal face.
Pseudoaperture (adj. pseudoaperturate) (Thanikaimoni, 1980)

A thinning of the exine which, although superficially resembling an aperture, is
not associated with a thickening of the intine and is presumed not to function as an exitus.
See also: pseudocolpus, pseudopore.
Pseudocolpus (pl. pseudocolpi, adj. pseudocolpate) (Iversen and Troels-Smith,

1950)
A colpus-like pseudoaperture. Example: Myosotis (Boraginaceae). See
also: heterocolpate.
Pseudopore (pl. pseudopores, adj. pseudoporate) (Iversen and Troels-Smith, 1950)

A pore-like pseudoaperture. Comment: The term has also been used for the
leptoma of certain coniferous pollen.
Pseudosaccus (pl. pseudosacci, adj. pseudosaccate) (Grebe, 1971)

An extensive, saccus-like separation in the wall of a spore resembling a saccus,
but lacking the characteristic alveolate infrastructure. Example: Grandispora spinosa. See
also: camera, saccus.
Proximal face (Erdtman, 1952)

That part of a palynomorph which faces towards the centre of the tetrad, between
equator and proximal pole. Antonym: distal face.
Psilate (adj.) (Wodehouse, 1928)
Describing a pollen or spore with a smooth surface.
Ptychotreme (adj.) (Erdtman and Straka, 1961)
With apertures situated in invaginations of the outline, when seen in polar view.
Comment: The term is not recommended because it is based upon the suffix-treme.
Punctum (pl. puncta, adj. punctate) (Erdtman, 1952)

A rounded or elongate tectal perforation, less than 1 μm in length or diameter.
See also: foveola, scrobiculus, tectum perforatum.
Quasisaccus (pl. quasisacci, adj. quasisaccate) (Meyen, 1988)

Synonym of protosaccus.
Quasitectate (adj.) (Balme, 1988)

A spore exine in which the outer and inner, more or less homogeneous, layers
are separated by a clearly defined mesexinous layer of discontinuous columellate elements,
simulating those that characterise many angiosperm pollen.
Radial
A general term describing features radiating from a centre. Applied in
palynology to the region of a spore beyond the ends of the laesurae.
Radially symmetric (adj.) (Nilsson and Muller, 1978)

Describing a pollen grain or spore with two or more vertical planes of
symmetry, but, if only two such planes are present, then their axes are of equal length.
Radius
A general term, used in palynology for a laesura in trilete spores.
Psilolophate (adj.) (Wodehouse, 1935)
Describing a lophate pollen grain which lacks spines. See
also: echinolophate, lophate.
Rectimurate (adj.) (Erdtman, 1952)
Describing a pollen grain or spore with more or less straight muri.
Reticuloid (adj.) (Moar, 1993)

With bacula arranged in a more or less reticulate pattern.
Synonym of retipilate.
Reticulum (pl. reticula, adj. reticulate) (Praglowski and Punt, 1973)

A network-like pattern consisting of lumina or other spaces wider than 1 μm
bordered by elements narrower than the lumina. See also: microreticum.
Retipilate (adj.) (Erdtman, 1952)

Describing a reticulum formed by rows of pila instead of muri. Example:
Callitriche (Callitrichaceae).
Retusoid (adj.) (Traverse, 1988)

Describing a spore with prominent contact areas and curvaturae. Example:
Retusotriletes.
Rhomboidal tetrad (Erdtman, 1945b)

A uniplanar tetrad in which the proximal faces of two individual members are in
direct contact and the remaining two are separated, giving a rhomboidal outline to the
tetrad. Example: Epipactis palustris (Orchidaceae).
Rimula (pl. rimulae, adj. rimulate) (Pflug, 1953)

The sub-equatorial aperture that encircles pollen grains of the Classopolles group.
Example: Corollina (Classopollis). Comment: Potonié (1934) used rimula to refer to short
colpi, but the term is no longer used in this sense.
Ruga (pl. rugae, adj. rugate) (Potonié, 1934, emend. Erdtman, 1945a)
Synonym of colpus.
Rupate (adj.) (n. rupus, pl. rupi) (Erdtman, 1952)
Synonym of loxoaperturate.
S-type tetrad (Moar, 1993)

A tetrad in which only one member is fully developed. Example: Leucopogon
fasciculatus (Epacridaceae). See also: A-type tetrad, T-type tetrad.
Saccoid (adj.) (Brugman, 1983)

A saccus-like expansion of the exine with a complex infrastructure consisting of
a three dimensional network of sexine elements, extending to and fused with the nexine.
See also: camera, protosaccus, pseudosaccus, saccus.
Saccus (pl. sacci, adj. saccate) (Erdtman, 1952)

A saccus formed by an expansion of the exine of a pollen grain and at least
partly filled with an alveolate infrastructure. See also: bisaccate, camera,
protosaccus, pseudosaccus.
Scabrate (adj.) (sing. scabra, pl. scabrae) (Iversen and Troels-Smith, 1950) Describing
elements of ornamentation, of any shape, smaller than 1 μm in all directions. Examples:
Quercus (Fagaceae), Artemisia (Compositae). Comment: Ornamentation elements larger
than 1 μm are described according to their shape, for example, baculum, clava, gemma,
verruca. See also: granulum.
Sclerine (Erdtman, 1952)

A term encompassing both exine and perine that can be used whether a perine is
present or not; sporoderm excluding the intine. See also: sculptine.
Scrobiculus (pl. scrobiculi, adj. scrobiculate) (Potonié, 1934)

Synonym of punctum.
Rugulate (adj., n. rugula, pl. rugulae) (Iversen and Troels-Smith, 1950) Describing a
type of ornamentation consisting of elongated sexine elements more than 1 μm long,
arranged in an irregular pattern that is intermediate between striate and reticulate.
Examples: Sedum (Crassulaceae), Ulmus (Ulmaceae), Camptotriletes corrugatus.
Sculptural density (Balme, 1988)

The estimated number of sculptural elements in an area of 100 μm2 of the surface
of the exine.
Sculpture (Kuprianova, 1948)

Orthographical variant of sculpturing.
Sculpturing (adj. sculptured) (Potonié, 1934)

The surface relief, or topography, of a pollen grain or spore. Comment:
Praglowski (1975) provided a circumscription of this term which would restrict its
application to tectate pollen grains. See also: pattern, ornamentation, structure.
Seed-megaspore
A large, functional megaspore associated with three small, presumably aborted
spores present in some fossil lycopsids. Example: Cystosporites. Semi-erect (adj.)
(Reitsma, 1970) Synonym of subprolate.
Semitectum (adj. semitectate) (Fægri and Iversen, 1964)

A partially discontinuous tectum in which the tectal perforations are equal to or
wider than the muri and usually larger than 1 μm in diameter. Antonym: eutectum. See
also: tectum.
Semitransverse (adj.) (Reitsma, 1970)

Synonym of suboblate.
Sexine (Erdtman, 1952)

The outer, sculptured layer of the exine, which lies above the nexine. Antonym:
nexine. Comment: Although the term ectexine as originally defined (sensu Erdtman,
1943) was more or less synonymous with sexine, it no longer is because ectexine is now
invariably used in the redefined sense of Fægri (1956). Unlike ectexine, sexine does not
include the foot layer. Sexine and nexine are distinguished on purely morphological
criteria, whereas ectexine and endexine differ in their staining properties. The two sets of
terms are therefore suited for slightly different applications. See also: ectexine.
Sexine 1, 2, 3, 4, 5 (Reitsma, 1970)

A system of sexine stratification in which sexine 1 is the innermost and sexine 5
in the outermost layer of the sexine. Comment: Usually the sexine consists of 3 layers
(sexine 1 = columellae; sexine 2 = tectum; sexine 3 = sclupture elements).
Sculptine (Erdtman, 1948)
A term encompassing both exine and perine (sclerine) but excludes the nexine
and so provides a neutral term for the sculptured layer when there is doubt whether the
pattern belongs to the exine or the perine. See also: sclerine.
Shape classes (pl.) (Erdtman, 1943)
Categories of pollen and spore shape based on the relations between polar axis
(P) and equatorial diameter (E). See also: oblate, oblate spheroidal, P/E ratio,
peroblate, perprolate, prolate, prolate spheroidal, suboblate, subprolate.
Simplibaculate (adj.) (Erdtman, 1952) Synonym of

simplicolumellate.
Simplicolumellate (adj.) (Reitsma, 1970)

With a single row of columellae under each murus. Example: Viburnum opulus
(Caprifoliaceae). See also: duplicolumellate, pluricolumellate.
Sinu-aperturate (adj.) (Erdtman, 1952)

Describing a pollen grain in which the equatorial apertures are situated in the
middle of concave sides when seen in polar view. Comment: The term belongs to the
system of ambs defined by Erdtman. See also: amb, angulaperturate, planaperturate.
Sole (APLF, 1975)

Synonym of foot layer.
Special wall (Beer, 1911) Pollen

mother-cell wall
Spheroidal (Erdtman, 1943)

Describing the shape of a pollen grain or spore in which the polar axis and the
equatorial diameter are approximately equal. Comment: This term belongs to the system of
shape classes suggested by Erdtman (1943), and extended in 1952), based on the
measurements of the polar axis (P) and equatorial diameter (E). In this system spheroidal is
defined as a P/E ratio of 0.88–1.14. See also: oblate spheroidal, prolate spheroidal.
Spina (pl. spinae) (Potonié, 1934)

Synonym of Spine.
Spinule (adj. spinulose) (Erdtman, 1952) Small spines, less than 3 μm in length. Comment:
The size distinction made by Erdtman between spinules and spines is not consistent with
other size criteria used in palynology. See also: spine.
Spiraperturate (adj.) (Erdtman, 1952)

Describing a pollen grain with one or more spiral apertures. Example: Eriocaulon
aquaticum (Eriocaulaceae). Comment: In the pollen classes of Iversen and Troels-Smith
(1950), spiraperturate pollen was included in the syncolpate class.
Spore
A general term for the usually microscopic, unicellular, asexual or sexual
reproductive units of cryptogams and fungi (Jackson, 1928). See also: pollen,
microspore.
Sporoderm (Bischoff, 1833)

The entire wall of a pollen grain or spore.
Sporomorph (Erdtman, 1947)

A general term for spore-like palynomorphs.
Sporopollenin (Zetsche and Vicari, 1931)

The name given to the acetolysis resistant biopolymers which make up most of
the material of the exine.
Square tetrad (Erdtman, 1945b)

Synonym of tetragonal tetrad.
Spine (adj. spiny/spinose) (Erdtman, 1952)

A general word, applied in palynology to long and tapering pointed elements,
exceeding 1 μm. Comment: In Erdtman's (1952) definition spines were defined as more
than 3 μm long (in contrast to spinules which were shorter than this) but since the
maximum size of features that may be prefixed with micro- is 1 μm the present definition
provides consistency with other terms. See also: echinate.
Stenopalynous (adj.) (Erdtman, 1952)
Describing plant taxa characterized by only a slight variation in their
palynomorphs. Antonym: eurypalynous.
Stephano-
Synonym of zono-. Comment: Fægri and Iversen (1950) used the prefix (as
for example, in stephanocolpate, stephanocolporate, stephanoporate) in their
classification of pollen types.
Stratum (APLF, 1975)
A subdivision of a major layer of the sporoderm.
Striae (pl.) (sing. stria) (Fægri and Iversen, 1950)

Grooves between elongated sculpturing elements.
Striate (adj.) (Iversen and Troels-Smith, 1950)
A general descriptive term applied in palynology to elongated, generally parallel
elements separated by grooves. Comment: The positive elements of striate ornamentation
may also be referred to as muri.
Striato-reticulate (adj.) (Erdtman, 1952)

Describing a pattern in which parallel or subparallel muri are cross-linked to
form a reticulum in the grooves. The connections between the muri lie on a single level or
different levels. Example: Gentiana pneumonathe (Gentianaceae).
Structure (adj. structurate, structured) (Potonié, 1934)

The internal construction of the pollen or spore wall. See
also: sculpturing, pattern, ornamentation.
Sub-A prefix for under, or less than.
Subechinate (Wodehouse, 1928)

Synonym of microechinate.
Suberect (adj.) (Reitsma, 1970)

Synonym of prolate spheroidal.
Subisopolar (adj.) (Walker and Doyle, 1975)

Describing a pollen grain or spore in which the proximal and distal faces are
slightly different. Examples: one face is convex and the other is less convex (Banksia,
Proteaceae); on one face viscin threads are connected (Oenothera, Onagraceae).
Sub-layer
Synonym of stratum.
Suboblate, (adj.) (Erdtman, 1952)

polar axis and the equatorial diameter is 0.75–0.88. See also: oblate, oblate spheroidal,
P/E ratio, peroblate, perprolate, prolate, prolate spheroidal, shape classes,
spheroidal, subprolate, subspheroidal.
Subprolate (adj.) (Erdtman, 1952)
polar axis and the equatorial diameter is 1.14–1.33. See also: oblate, oblate
spheroidal, P/E ratio, peroblate, perprolate, prolate, prolate spheroidal,
shape classes, spheroidal, suboblate, subspheroidal.
Subspheroidal (adj.) (Erdtman, 1952)

polar axis and the equatorial diameter is 0.75–1.33. Comment: This shape class includes
suboblate, oblate spheroidal, prolate spheroidal and subprolate. See also:
oblate, oblate spheroidal, peroblate, P/E ratio, perprolate, prolate, prolate
spheroidal, shape classes, spheroidal, suboblate, subprolate.
Subtransverse (adj.) (Reitsma, 1970) Synonym of

oblate spheroidal.
Subturma (pl. subturmae) (Potonié, 1956)

A group of form-genera of fossil spores or pollen in the turma-system of
Potonié. See also: anteturma, infraturma, turma.
Successiform (Van Campo, 1967)

Referring to a phyletic series of pollen types with increasing numbers of
apertures, ranging from tricolpate to pantocolpate and pantoporate.
Sulculus (pl. sulculi, adj. sulculate) (Erdtman, 1952)

An elongated latitudinal ectoaperture not situated at a pole. See also:
sulcus.
Sulcus (pl. sulci, adj. sulcate) (Erdtman, 1952)

An elongated latitudinal ectoaperture situated at the distal or proximal pole of a
pollen grain. Comment: A sulcus has the same shape as a colpus, but differs in
orientation. Sulci are essentially latitudinal apertures whereas colpi are essentially
longitudinal apertures. Sulci may be distal (anasulcate), proximal (catasulcate) or
extend right around the grain (zonasulcate). See also: ana-, colpus, zona-.
Supra-
A prefix for above. Comment: In palynology this prefix is mostly used for
features on top of the tectum, as for example in suprareticulate, suprarugulate,
suprastriate.
Supratectal (adj.) (Erdtman, 1969)

Indicating the position of features, such as spines, on the top of the tectum.
Suture (Potonié and Kremp, 1955)

Synonym of commissure. See
also: laesura.
Sylvestris-type (Rudolph, 1935)

Bisaccate pollen grains in which the outline of the sacci in polar view is
discontinuous with the outline of the corpus so that the grains seem to consist of three
distinct, more or less oval parts. Examples: Pinus sylvestris, Abies (Pinaceae). See also:
diploxylonoid, haploxylonoid, haploxylon-type.
Syn-A prefix indicating the fusion or anastomosis of features.
Syncolp(or)ate (adj.) (sensu Erdtman, 1952)

Describing a pollen grain with two or more simple (or compound) colpi the ends
of which anastomose at the pole. Example: Primula farinosa (Primulaceae). Comment: The
term was used in a wider sense by Iversen and Troels-Smith (1950) for other forms with
fused apertures such as spiraperturate and parasyncolp(or)ate.
Synrugoidate (adj.) (Erdtman, 1952; Jalan and Kapil, 1964)

Describing a pollen grain with six colpi of which three are long and meeting at
one pole and three are short and not meeting at either pole. Example: Schisandra
grandiflora (Schisandraceae).
T-shaped tetrad (Walker and Doyle, 1975)

A uniplanar tetrad in which two of the members are perpendicular to the
other two so that tetrad has the shape of the letter “T”. Example: Typha spp. (Typhaceae).
See also: tetrad.
T-type tetrad (Moar, 1993)

A tetrad in which all four members are fully developed. Example: Pentachondra
pumila (Epacridaceae). See also: A-type tetrad, S-type tetrad.
Tectum (pl. tecta, adj. tectate) (Iversen and Troels-Smith, 1950)

The layer of sexine, which forms a roof over the columellae, granules or other
infratectal elements. See also: eutectum, semitectum, tegillum.
Tectum perforatum (adj. tectate perforate) (Iversen and Troels-Smith, 1950)

A tectum with perforations smaller than 1 μm in diameter. See also:
punctum.
Tectum imperforatum (adj. tectate imperforate) (Walker and Doyle, 1975)

With a continuous tectum, without perforations.
Synonym of eutectum.
Tectum solidum (Fægri and Iversen, 1975)

Synonym of eutectum.
Tegillum (pl. tegilla, adj. tegillate) (Erdtman, 1952)

Synonym of tectum.
Taenia (pl. taeniae, adj. taeniate) (Leschik, 1956)

One or more strap-like, more or less parallel strips of exine on the proximal
and/or distal sides of the corpus of certain fossil gymnospermous pollen grains. Examples:
Striatites, Vittatina. See also: polyplicate.
Tenui- (Erdtman, 1952)
A prefix for thin. Comment: Erdtman often used this prefix in combination with
other terms (e.g. tenuimarginate) but thinness is subjective; it is preferable to use alternative
descriptions.
Tenuitas (pl. tenuitates) (Potonié, 1934)

A general term for a thinning, that has been applied to many different situations
in palynology. Comment: The term was used by Potonié to describe endoapertures, by
Potonié and Kremp (1955) for sulci and has sometimes been used for the distal thin parts in
the exine of the Circumpolles group (Corollina).
Tetrad mark (Couper and Grebe, 1961)
The monolete or trilete mark on the proximal face of a spore or, more rarely, a
pollen grain. See also: trilete mark, Y-mark.
Tetrad stage
The period during post-meiotic development when the four microspores or
megaspores are united by the presence of a temporary special wall. Comment: The tetrad
stage ends at the start of the free spore stage, when the special cell wall is reabsorbed.
Tetragonal tetrad (Grebe, 1971)

A uniplanar tetrad in which all four members are in contact at the centre of the
tetrad so that, in the correct orientation, the adjacent walls form a cross. Example: Uvariastrum
hexaloboides (Annonaceae). See also: tetrad.
Tetrahedral tetrad (Grebe, 1971)

A multiplanar tetrad in which each member is in contact with three others, so that
the centres of the grains define a tetrahedron. Example: Erica (Ericaceae). See also:
decussate tetrad, tetrad.
Tilia structure (Praglowski, 1971)

A sexine structure describing a pertacte tectum provided with funnel-like
concavities of which the bottoms coalesce with a single underlying columella, to form one unit.
Torus (pl. tori) (Thomson and Pflug, 1953)

An arcuate invagination or protuberance of the exine more or less paralleling the
laesura of a Spore in the interradial area. See also: kyrtome, labrum.
Tetrad
A general term for a group of four united pollen grains or spores, either as a
dispersal unit or as a developmental stage. Comment: Tetrads may be uniplanar, with all
members lying in the same plane (for example, linear, rhomboidal, tetragonal and
T-shaped tetrads) or multiplanar, with members in more than one plane (for example,
decussate or tetrahedral tetrads). See also: dispersalunit, monad, dyad.
Transversal furrow (Iversen and Troels-Smith, 1950)

Synonym of endocolpus.
Trema (pl. tremata) (Erdtman and Straka, 1961)

Synonym of aperture. See
also: -treme.
-treme (Erdtman and Straka, 1961)

A suffix synonymous with aperture. Comment: The -treme system of aperture
classes suggested by Erdtman and Straka was intended to replace that of Iversen and
Troels-Smith (1950). The system included atreme, monotreme, ditreme, tritreme,
tetratreme, pentatreme, hexatreme, polytreme, anomotreme, pleotreme and stephanotreme.
Tri-A prefix for three.
Trichotomocolpate (adj.) (Erdtman, 1945a)

Describing a pollen grain with a three-armed colpus. Example: Trapa natans
(Trapaceae).
Trichotomosulcate (adj.) (Erdtman, 1952)

Describing a pollen grain with a three-armed sulcus. Example: Elaeis guineensis
(Palmae).
Tricolpate, tricolporate, triporate (adj.) (Iversen and Troels-Smith, 1950) Describing

pollen grains with three ectocolpi, three compound apertures or three pores. Comment: The
terms belong to the system of pollen classes introduced by Iversen and Troels-Smith
(1950). See also: monoaperturate, pantoaperturate, zonoaperturate.
Transverse (adj.) (Reitsma, 1970)
Synonym of oblate. Comment: Transverse has also been used to indicate the
orientation of latitudinal features. It is also commonly used to indicate a section cut
through the equatorial plane of a pollen grain.
Trilete (adj.) (Erdtman, 1943)

Describing a spore with three laesurae, thus showing a trilete mark.
Example: Pteridium (Hypolepidaceae). See also: alete, laesura, monolete.
Trilete mark
The triradiate mark of a trilete spore.
Triprojectate (adj.) (Mtchedlishvilli, 1961)

Describing projectate pollen with three projecting arms on which the apertures
are situated, as for example in the Triprojectacites group of fossil pollen grains. Example:
Aquilapollenites.
Tryphine (Erdtman, 1969)

A material deposited on the surface of pollen grains by the breakdown of the
tapetum and differing from pollenkitt in that it contains membraneous components
derived from organelles. See also: pollenkitt.
Tuberculate (adj.)
A general term for beset with knobby projections or excrescenses (Jackson,
1928).
Tubulus (pl. tubuli) (Erdtman, 1952)

A general term for a small channel, applied in palynology to a channel through
the nexine.
Tula (pl. tulae, adj. tulate) (Jansonius and Pocock, 1969)

A sexinal inflation in gymnospermous pollen at the end of the axis of the distal
sulcus or leptoma. Example: Ovalipollis.
Tumescence (pl. tumescences) (Batten and Christopher, 1981)
A gradual increase in thickness of the wall layer(s) from a point in the equatorial
interradial region to the germinal aperture. Example: Megatriopollis. Comment: This term
is used in descriptions of the fossil Normapolles group.
Trifolium (pl. trifolia) (Potonié, 1956)
A three-bladed, proximal feature arising from the proximal pole of a
megaspore. Example: Capulitriletes. See also: acrolamella, gula.
Turma (pl. turmae) (Potonié, 1956)
An artificial suprageneric grouping of form-genera of fossil spores and pollen.
Comment: The following groups are recognised in the system: anteturma, turma,
subturma, and infraturma.
Turriform (Balme, 1988)

Biform sculptural elements consisting of a capitate basal portion surmounted by a
sharply contracted distal spine. Example: Dibolisporites.
Ubisch body (pl. ubisch bodies) (Rowley, 1963)

A distinctive, orbicular granule of sporopolleninsporopollenin produced by the
tapetum, particularly in plants with secretory tapeta. See also: orbicule.
Ulculus (pl. ulculi, adj. ulculate) (Walker and Doyle, 1975)

A rounded ectoaperture not situated at a pole. Example: Poaceae. See also:
ulcus.
Ulcus (pl. ulci, adj. ulcerate) (Erdtman, 1952)

A rounded ectoaperture situated at the distal or proximal pole of a pollen grain.
Examples: Sparganium (Sparganiaceae), Typha (Typhaceae). Comment: An ulcus has the
same shape as a pore. Ulci may be distal (ana-ulcerate) or proximal (cata-ulcerate).
See also: ana-, cata-, pore, ulculus.
Uniplanar tetrad (Walker and Doyle, 1975)

A tetrad in which the individual members lie more or less in one plane.
Urceolate (adj.) (Ferguson et al., 1983)
Describing a type of ornamentation consisting of urn-shaped elements situated on
the footlayer. Example: Pinanga aristata (Palmae).
Vallum (pl. valla) (Iversen and Troels-Smith, 1950)

Synonym of murus. Comment: Sometimes used to describe a single, broad,
raised feature in certain fossil spores. See also: lira.
Valva (pl. valvae, adj. valvate) (Potonié and Kremp, 1955)

Radial thickenings in the areas beyond the ends of the laesurae of trilete spores.
Example: Triquitrites tribullatus. Comment: The term auriculate (Potonié and Kremp,
1955) refers to an extremely valvate condition.
Velum (pl. vela, adj. velate) (Thomson and Pflug, 1953)

A feature of a monosaccate pollen grain in which the saccus is convoluted.
Example: Tsuga (Pinaceae).
Vermiculate (adj.) (Kosanke, 1950; Harris, 1955)

A general descriptive term used to describe winding features. Comment:
Vermiculate has been used by Kosanke (1950) and Harris (1955) to describe depressions
(fossulae). Others use the term for raised structures (muri) in rugulate pollen and
spores.
Verruca (pl. verrucae, adj. verrucate) (Iversen and Troels-Smith, 1950)

A wart-like sexine element, more than 1 μm wide, that is broader than it is high
and is not constricted at the base. Example: Plantago (Plantaginaceae).
Verrucose (adj.) (Erdtman, 1969)

Synonym of verrucate.
Vacuoles (Grebe, 1971)
Rounded to elongated spaces within an exinous structure, e.g. a cingulum or
zona. Example: Vallatisporites ciliares. Synonym of dissections. Comment: In botanical
sense, vacuole is a general term for a liquid filled cellular component (Jackson, 1928). In
Latin it means a hollow space, and in this sense this term is used.
Vestibulum (pl. vestibula, adj. vestibulate) (Potonié, 1934)

A separation between layers of the exine forming a cavity between the inner and
outer pores. For example: Betula (Betulaceae). See also: atrium, fastigium.
Viscin thread (pl. viscin threads)

A general botanical term (Jackson, 1928), applied in palynology for an acetolysis
resistant, sporopollenin thread arising from the exine of a pollen grain, usually from the
proximal surface. Example: Oenothera (Onagraceae).
Y-mark (Potonié, 1934)

Synonym of trilete mark.
Z-layer (Heslop-Harrison, 1979)

Synonym of exintine.
Zona (pl. zonae, adj. zonate) (Potonié and Kremp, 1955)
A thin outer structure of a spore that projects at the equator, but does not extend
over the distal face or proximal face. Example: Cirratriradites saturni. Comment: The term
is especially applied for many representatives of fossil spores. Example: Kraeuselisporites.
See also: cingulum, corona.
Zona- (adj. zonate) (Walker and Doyle, 1975)

Prefix indicating ring-like. Comment: The prefix zona- is used in conjunction
with a suffix indicating the type of aperture.
Zona-aperturate (adj.) (Walker and Doyle, 1975)

Describing a pollen grain with a ring-like aperture. Comment: The range of
ring-like apertures includes for example: anazonasulculus (1), a ring-like sulculus in the
distal hemisphere; catazonasulculus, a ring-like sulculus in the proximal hemisphere;
zonasulculus, a ring-like sulculus around the equator; and zonasulcus (2), a
meridional ring-like sulcus perpendicular to the equator.
Vesiculate (adj.) (Iversen and Troels-Smith, 1950)
Synonym of saccate.
Zonasulcus (Walker and Doyle, 1975)

A meridional ring-like sulcus perpendicular to the equator. Example: Laurelia
novaezelandiae (Monimiaceae). See also: meridionosulcus.
Zoni- (sensu Erdtman and Vishnu-Mittre, 1956)

Synonym of zono-.
Zoni- (sensu Walker and Doyle, 1975)

Prefix indicating a latitudinal orientation.
Zono- (Erdtman and Straka, 1961)

A prefix indicating features located equatorially. See
also: ana-, cata-, stephano-.
Zonoaperturate (adj.) (Erdtman and Straka, 1961)

Describing a pollen grain with apertures situated only at the equator. Comment:
Pollen classes with zonoaperturate pollen include, zonocolpate, zonocolporate,
zonoporate. Originally Erdtman and Vishnu-Mittre (1956) introduced the prefix zoni-,
but later Erdtman and Straka (1961) changed the ending of the prefix to zono-. See also:
stephano-.
Zonorate (adj.) (Erdtman, 1952)

Describing a pollen grain with a continuous endoaperture (os) around the
equator.
Synonym of endocingulum.
Zwischenkörper (pl.) (Fritzsche, 1837)

A lens-shaped body below the aperture of certain unacetolysed pollen grains.
Comment: The feature resembles an oncus but is treated as distinct because some pollen
grains have both features.
Zonasulculus (Walker and Doyle, 1975)
A ring-like sulculus around the equator. Example: Nymphaea violacea
(Nymphaeaceae). See also: meridionosulcus.
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Glossary of pollen and spore terminology

Received 28 June 2006; accepted 28 June 2006 Available online 28

Wim Punt, Stephen Blackmore, Siwert Nilsson a

into question the utility of the Group's methodology. During a

A-type tetrad (Moar, 1993)

Abporal lacuna (pl. abporal lacunae) (Wodehouse, 1928)

Acalymmate (adj.) (Van Campo and Guinet, 1961)

Acanthomamilla (Balme, 1988)

Acetolysis (Erdtman, 1960a)

Acolpate (adj.) (Moar, 1993)

Acrolamella (pl. acrolamellae, adj. acrolamellate) (Li and Batten, 1986)

Alete (adj.) (Erdtman, 1943)

Alveolate (adj.) (Van Campo, 1971)

Amb (Erdtman, 1952)

Ana-(Erdtman and Vishnu-Mittre, 1956)

Anazonasulculate (adj.) (Walker and Doyle, 1975)

Ancyrate (adj.) (Balme, 1988)

Angulaperturate (adj.) (Erdtman, 1952)

Anisopolar (adj.) (Erdtman, 1947)

Annulus (pl. annuli, adj. annulate)

Anteturma (pl. anteturmae) (Potonié, 1956)

Aperture (adj. aperturate) (Erdtman, 1947)

Apicalfeld (Beug, 1961)

Apiculate (adj.) (Fægri and Iversen, 1950)

Apiculate elements (Smith and Butterworth, 1967)

Apocolpial field (Punt et al., 1974)

Apocolpium (pl. apocolpia) (Erdtman, 1952)

Apocolpium index (Punt, 1976)

Arcus (pl. arcus, adj. arcuate) (Erdtman, 1947)

Areola (pl. areolae, adj. areolate) (Erdtman, 1947)

Aspidote (adj.) (Erdtman, 1952)

Aspis (pl. aspides, adj. aspidate) (Wodehouse, 1935)

Aspis channel (Engel, 1980)

Apoporium (pl. apoporia) (Erdtman, 1952)

Auricula (pl. auriculae, adj. auriculate) (Potonié and Kremp, 1955)

Baculum (pl. bacula, adj. baculate) (Potonié, 1934)

Bireticulate (adj.) (Bor, 1979)

Atectate (adj.) (Walker and Doyle, 1975)

Atrium (Thomson and Pflug, 1953)

Atrium (Punt, 1962)

Bisaccate (adj.) (Potonié and Kremp, 1954)

Breviaxy (Van Campo, 1966)

Bridge (Fægri and Iversen, 1950)

Brochus (pl. brochi, adj. brochate) (Erdtman, 1952)

Callose wall (Mangin, 1899) Part of the

Calymmate (adj.) (Van Campo and Guinet, 1961)

Cata- (Erdtman and Vishnu-Mittre, 1956)

Capillus (pl. capilli, adj. capillate) (Potonié and Kremp, 1955)

Cappa (pl. cappae) (Erdtman, 1957)

Cappula (pl. cappulae) (Erdtman, 1957)

Capsula (pl. capsulae, adj. capsulate) (Pocock, 1961a)

Cavea (pl. caveae, adj. caveate) (Skvarla and Larson, 1965)

Caverna (pl. cavernae) (Thomson and Pflug, 1953)

Cavium (Thomson and Pflug, 1953)

Cavum (pl. cava, adj. cavate) (Fægri and Iversen, 1989)

Cavus (sensu Skvarla and Turner, 1966)

Centrosymmetrical (adj.) (Straka, 1964)

Circumaperturate (adj.) (Straka, 1964)

Clava (pl. clavae, adj. clavate) (Iversen and Troels-Smith, 1950)

Coaperturate (adj.) (Beug, 1961)

Colpodiporate (adj.) (Selling, 1947)

Colpororate (adj.) (Moar, 1993)

Colpoid (Erdtman, 1952)